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ABSTRACT: The burrowing land crabs of the world are defined as an ecological group and the burrow or crabhole fauna1 community is recognized and discussed as such. Introductory remarks on terminology, relationship of the crabhole habitat to other habitat types, general physical nature of the crabhole, and the major ecological structure of the community as now known are presented. The remainder of the paper consists of two parts: 1) A list of all the species of borrowing land crabs of the world, including notations on distribution, recognition, and ecology. Twenty-four species in the genera Sesarma, Ocypode, Uca, Ucides, Gecarcoidea, Cardisoma, and Gecarcinus are given. 2) A list of all published records of arthropods found in crab burrows either associated with the crab as a burrow coinhabitant or having symbiotic relationships with it. The vast majority of these are insects, primarily mosquitoes, of which 140 species are noted. For each burrow associate or symbiont, the distribution, recorded crab host, type of relationship (specific, semispecific, transient or accidental) are given.

INTRODUCTION The present paper represents a literature survey to establish the present state of knowledge on the unique ecological relationship existing between burrowing land crabs and a variety of associated organisms. From our own field studies it is evident that there are many unrecorded species of arthropods occurring in crabholes and undescribed ecological phenomena to be discovered and analyzed. We hope that from this beginning other workers will recognize the land crab burrow as a special habitat and respond to the need for inquiring further into its natural history. Published data on land crabs consists primarily of species accounts and selected aspects of behavior and natural history; no broad coverage of the basic ecology of any species exists. Likewise, with regard to the burrow associates, no general ecological treatment is available, only taxonomic notes and fragmentary collecting data. We are presently engaged in a project to study the biology of land crabs and their burrow associates (Hogue and Bright, 1969). One preliminary field survey in Kenya, East Africa (Hogue and Bright, 1971) has been reported. Field studies in Costa Rica, Baja California, Pacific mainland
IDonald B. Bright, Associate Professor of Biology, lerton, California 9263 1, California Natural State Colege, FulHistory Museum

2Charles L. Hogue, Senior Curator of Entomology, of Los Angeles County, Los Angeles, California 90007.

and crustaceans in the Department of Biology. Ocypodidae. Entomological materials are deposited in the Los Angeles County Museum of Natural History. Panama. DEFINITIONS Since we make use of terms and concepts originally devised for very different community types. certain species only Uca. Burrowing land crab: This term refers to a group of tropical species which dig well-defined burrows above the normal flooding and flushing action of the tides. Though we consider these outside our specified limits with burrowing land crabs (and they are omitted from the section on burrowing land crabs). certain species only Several species of land dwelling lobsters (e. Thalassina). 220 Mexico. all species Cardisoma. It is presently possible to evaluate only the gross community structure of arthropod symbionts and burrow associates of land crabs in terms of the general levels of interrelationships and the most conspicuous variations in niches displayed by the arthropod fraction. By niche we mean the total ecological role a species plays in the community. These species belong to the families Gecarcinidae. All specimens and data collected on this project are given the code LCBA (Land Crab Burrow Associates). we mention them in our survey of associates because they occur sympatrically with burrowing land crabs and confusion frequently arises in identifying the true owner of a burrow (Scharff and Tweedie. California State College. both in regard to its habitual location of occurrence (place niche-microhabitat) . Western Caribbean (Islas San And& and Providencia). Peru. 1942). all species Grapsidae (only 1 of 26 genera in the family) Sesarma. crayfish (Procambarus. Cambarus) and freshwater crabs (Sudanonautes) also construct burrows supporting an associated fauna. Thus our attention will be directed to the following taxa: Gecarcinidae (all 3 genera of the family) Gecarcoidea.. we find it necessary to define certain of our present usages: Crabhole and crab burrow: Used synonymously.g. Ecuador.2 CONTRIBUTIONSIN SCIENCE No. Fullerton. certain species only Ocypodidae (3 of 4 genera in the family) Ocypode. certain species only Ucides. Ecological structure of crabhole community: All of the organisms found in the crabhole and associated ecologically with land crabs we refer to as the crabhole community. The family Coenobitidae is excluded since the terrestrial hermit crabs are non-burrowers. and Grapsidae. and Australia will be reported upon in forthcoming papers. all species Gecarcinus.

Major Examples Immature mosquitoes Diving beetles (Bidessus) Cyclops Burrow chambers and surface of burrow water SYMBIOSIS Parasitism Resting. Adult mosquitoes Adult biting gnats (Culicoides) Mite (Laelaps cancer) Gill chamber of Gecarcinus la teralis Peribuccal cavity and renal grooves of Gecarcinus ruricola Attaching and feeding Attaching to host and feeding on food debris Commensalism Drosophila carcinoplzila . refers to those species not only tied together by these factors but which also depend directly upon some form of intimate (often or usually also involving prolonged physical contact) interaction with one other member of the community. etc.e. etc. mating.. feeding.1972 BURROWING LAND CRABS OF THE WORLD 3 and its inter-dependency with other members of the community (functional niche). Specific (or obligatory) The species is narrowly adapted to conditions in the crabhole or lives symbiotically with the host or other community member. parasitism or commensalism (no example of mutualism yet having been found). symbiosis. The extent to which a particular organism is dependent on the crabhole as a suitable habitat and to which it is an obligate member of the crabhole community may be further classed: I. A summary of the general ecological structure of the arthropod fraction of the crabhole community is given in Table 1. The second. breeding. Such species have specific adaptations to the physical and biological stresses encountered in the burrow (the precise nature of and adaptive significance of which are TABLE I SUMMARY OF THE KNOWN ECOLOGICAL STRUCTURE OF THE CRABHOLE COMMUNITY (ARTHROPOD FRACTION) Level of Interspecies Reaction ASSOCIATION Niche Place Burrow water Functional Developing. is shown by the assemblage of species whose common occurrence is dictated directly by the spectrum of indigenous limiting factors (physical and biotic) encountered in the crabhole. i. simple nssociation. Within the crabhole community we find two general levels of interspecific reaction : The first of these.

open sea. III. Examples occur among the following genera : Deinocerites.5-3. These four categories of habitat dependence. II. these mosquitoes require certain aquatic plants to which they attach with special respiratory structures for extracting vascular oxygen. Aedes (Cancraedes and Geoskusea). etc. An example is the mosquito Aedes (Skuseu) pembaensis which habitually breeds in crabholes but also develops commonly in various other types of coastal ground water accumulations (pools. reduced salt absorbing organs. Some species. For example. crabhole water dwellers have wide osmoregulatory capacities. wind blown adults.) or directly from rainfall. impermeable cuticle. etc. Transient (or facultative) The species usually inhabits other sites but may take up temporary residence or breed in the crabhole because of its similarity or proximity to the normal habitat. Aedes (Stegomyia) polynesiensis. larvae flushed from ground pools during heavy rains. are provisional. Examples are the many mosquitoes occurring along the seashore such as Aedes (Neomacleaya) panayensis and Aedes (Ochlerotatus) taeniorhynchus.4 CONTRIBUTIONSIN SCIENCE No. ponds.g. While there is considerable specific variation. Culex (Cu2ex) annulirostris. 220 presently little known. The diameter generally is equivalent to the carapace width of the host crab and the depth is determined by the level of the water table. in general the land crab burrow is a gently sloping or near vertical tubular excavation ranging from depths of 1. adults of Mansonia mosquitoes are sometimes found in crabholes but do not develop from larvae and pupae living in the burrow. the bottom of the burrow is filled with water derived indirectly from ground seepage from nearby sources (streams. Accidental The species is adapted to another habitat and only rarely occurs in and about crabholes for some anomalous reason (e.4 m.).). Unfortunately. of course. Like estuarine organisms in general. and indiscriminate breeders. in genera such as Gecarcinus. Semispecific (or semiobligatory) The species usually inhabits the crabhole. for no species do we yet know the complete story of its functional niche in the crabhole community.. are so well adapted to terres- . such as prolonged developmental period. estuaries. GENERAL PHYSICAL NATURE OF THE HABITAT IV. which are able to accommodate to such changes physiologically. These plants live only in open freshwater pools and ponds. Normally. being adapted at least in part to certain of its conditions but survives well in other habitats. etc. Thus the water may vary considerably in solute concentrations even from day to day or hour to hour. insular treehole breeders. swamps and even in artificial containers). and Drosophila.

Ocypode occidentalis 8. In listing the various species we have assumed that the published reports on burrow ownership are correct. Burrows also are often found along large rivers far inland where the fresh water affords the crab’ hydrobiotic needs. the species of Cardisoma and Sesarmcr..1972 BURROWINGLAND CRABSOF THE WORLD 5 triality that their burrows often are for physical protection only and penetration to the water table to maintain a supply of water for physiological functions is not necessary. that are likely to be additional hosts. No attempt has been made to provide a complete synonymy. The species we define as land crabs are enumerated below followed by a synopsis of important information on each species. is depauperate. Where there is considerable taxonomic confusion a note to clarify our usage has been included in the species accounts. Ocypode quadrutu 9. specific localities and/ or habitats. based on our field experience. Ocypode gaudichuudii 7. This is particularly true for distributional patterns in the Indo-Pacific. The burrows are located most often in compact alluvial soil well above high tide lines but still close enough to the sea to permit migration for spawning and close enough to a ground water source to maintain a reservoir. Sesurmu (Chiromun tes) africunurn 4. (See Tweedie ( 1950) for a discussion of the problems associated with Cardisoma. Sesarma (Sesurrnu) meinerti 3. e. In several of the species accounts there is question regarding reliable taxonomic and zoogeographic data. The community of these shallow burrows. Sesavvna (Sesarma) sulcutum 2.) Correspondence with a number of workers indicates that revisions are in preparation for Sesarma. Where authors were unable to identify the crustacean but provide descriptions. Sesurma (Holometopus) eulimene Family: Ocypodidae 6. Ocypode ceratophthulmu . Family: Grapsidae 1. Sesurmu (Holometopus) ortmunni 5. These works should aid future studies on the distribution of land crabs and their burrow associates. Uca and Gecarcinus. we have sometimes made a provisional determination. lacking the aquatic fraction. It is hoped that in future accounts authors will attempt to determine hosts specifically and include additional remarks on general ecology of the crab and its associates.g. s LAND CRABS OF THE WORLD The following genera and species accounts are for those burrowing land crabs listed as hosts in the arthropod portion of this paper or those.

Ucides occidentalis No. D-distribution. Crosnier. Uca tangeri 13. Some do not construct burrows but live under debris (rocks and roots). Note: Considerable taxonomic confusion prevails in this taxon. Most individuals are solitary. epistome well defined. 220 Family: Gecarcinidae humei guanhumi crassum armatum carnifex hirtipes planatus ruricola quadratus lateralis 15. mud flats.T) . Rathbun. 1880 (D).) . physiology. 1914 (T).e. Crane. Macnae. Gecarcoidea 16.T) . oval and occupy only slightly less than half of the anterior border of the carapace. Cardisoma 17. 22. B-biology. gravelly mud along lagoons and mangroves. 1940 (D. 1817 CHARACTERS: Carapace squarish. chelipeds thick. Uca subcylindrica 12. habitat. 21. 1947 (B. G-general). Ucides cordatus 14. banks of drying streams. mud-gravel areas extending from the surface down 1 m to the water table. 1955 (D.D. Miers. 1965 (D. life history. In some areas young live in the same burrow with an adult. and subequal in male. and third pair of legs longest. habits. 23. Cardisoma 18. 24.T. These burrows are in muddy.6 CONTRIBUTIONSIN SCIENCE 10. sides generally straight and parallel.T) . antennules transverse. particularly the validity of the subgeneric groupings. 1934 (D) . orbits of eye deep. Campbell. Many synonymies are suspected 1The application of each reference is indicated by a symbol following (T-taxonomy. DISTRIBUTION: Tropical and subtropical coastal areas of the world. Uca pugilator 11.D) . Tweedie. REFERENCE+: Bott.T). i. Cardisoma Gecarcinus Gecarcinus Gecarcinus Gecarcinus Family GRAPSIDAE Genus Sesarma Say. HABITS: Generally these have well defined burrows which are similar to those of most Cardisoma juveniles.. . Gordon. 1966 (B. HABITAT: Coastal marshes. Cardisoma 19. Cardisoma 20. 1967 (D.T). Tesch. 1917 (D.

Crane.T. Madagascar. Sesarma (Sesarma) sulcatum Smith. across Indo-Pacific to s Australia (Cooktown) and north to the Philippines. lower portion of male chelae cream yellow.. The burrows are deep and usually extend to the water table.T) . some species determinations are questionable. Individuals are solitary. COMMON NAMES: Mangrove crab.. 1870 COLOR: Carapace and legs dark brownish gray.. HABITS: They construct straight or slightly sloped burrows or live under debris characteristic of the habitat. 1947 (B. Sesarma (Sesarma) rneinerti de Man. and chelipeds a striking brilliant red. Mauritius. Salicornia (Baja California) and the roots and pneumatophores of mangroves. They are active throughout most of the day (except in drier areas) feeding primarily on plant materials.g.D. Cott (1930) gives a good account of this plus a consideration of the theory of warning colors.g. Mozambique. anterior and lateral margin bordered with orange to light yellow. Speckled crab. e. 1887 COLOR: Carapace black to gray or purple to deep violet.. e.1972 BURROWINGLAND CRABSOF THE WORLD because of the inordinate number of species. east coast of Africa (south to Port St..g. 1955 (D. 1966 (G). 1960 (D). and on the banks of streams and mangroves. Wright. remaining at the mouth of the burrow. HABITAT: Gravelly mud along lagoon shores. e. Sueda. John’ ) . 1966). 1. Marsh crab. They are often sympatric also with Goniopsis pulchra. Often the mouth of the burrow has a hood built of mud excavated while enlarging the tunnel or cleaning out. Ucides occidentalis and Uca spp.) . They apparently feed primarily on plant material. In drier habitats they commonly use burrows (occupied and unoccupied) of other crabs. chelae in females cream with a few maroon striations. Rhizophora and Avicennia (Costa Rica and Panama). three to six individuals under a rock (Wright. DISTRIBUTION: Andamans and Madras. Baja California to southern Panama). 115 in the Indo-Pacific alone. relatively hard mud. e. and only leave to forage at night. HABITS: Burrows are well developed and most common in areas where there is dry. tidal marshes. underside a dirty yellow. REFERENCES: Bott. Campbell (1967) and Crosnier (1965) give recent accounts dealing with these problems. 2. but also act as scavengers where . muddy banks associated with estuaries and mangroves. Garth. Because of the above. Cardisoma crusswn. females with conspicuous yellow line across front. drier. e.g. These crabs are retiring. but in areas where burrow structure is not well developed they tend to occur in groups.g. Some spend considerable time climbing the branches of marsh plants. HABITAT: Sandy-clay areas and higher. DISTRIBUTION: Pacific coast of the Americas (San Ignacio Lagoon.

and often associated with the pneumatophores of mangroves. Typically. 197 1 (D) .T). 1965 (D. when threatened.T) . Burrows generally well developed. 1954 (G) . Chace. HABITS: Constructs shallow burrows among exposed pneumatophores of the nangrove. Macnae and Kalk. 1954 (G) . Millard and Harrison. REFERENCE: Rathbun. COMMON NAME : None recorded.8 CONTRIBUTIONSIN SCIENCE No. 1837 COLOR: Carapace reddish brown.3-. 1953 (D. salt marshes. Crosnier. Sesarma (Chiromantes) africanurn H.D) . 1966 (B. HABITS: Occurs primarily in dense. chelae a dull to bright orange. . 1947 (D. 1965 COLOR: Carapace greenish brown and heavily calcified. Angola. Avicennia. COMMON NAME: None recorded. they hide under debris and roots. Sesarma (Holometopus) ortmanni Crosnier. 4. with soft mud. since they have no burrows or only very small ones. climbing over vegetation. Millard and Harrison. HABITAT: Muddy soil along the margins of mangroves. underside a dirty white. 192 1 (D. and mouths of rivers. also Barbados (?) . Millard and Harrison ( 1954) indicate extensive distribution of this species in Richards Bay. Crosnier.T) . South Africa.T) . Macnae and Kalk. but shallow. transverse patches of stiff hairs over carapace and limbs. Madagascar. Hogue & Bright. 1969 (B. Cott. COMMON NAME : Hairy lagoon crab. 1900 (D. DISTRIBUTION : Senegal to Benguela. in areas along the mangrove margins where there is deep. DISTRIBUTION: East coast of Africa from Malindi to Durban. 1930 (G) . HABITAT: Mangroves.T) . REFERENCES: Crosnier. Macnae. soft mud covered at least once per day by tidal flux. distal portion of chelae violet-red in color. Milne Edwards.D) . These burrows are not known to intersect.D) .T) . REFERENCES: Barnard. 1950 (D. HABITAT: Mud areas of salt marshes and mangroves. COMMON NAME : Marsh crab? REFERENCES: Alcock. 3. Sesarma (Holometopus) eulimene de Man. Juveniles and adults are conspicuous. 1965 (D. they construct shallow (. chelae of male bright orange-red. well shaded areas of mangroves. DISTRIBUTION: East coast of Africa. 1965 (D.T) . HABITS: Poorly known. 1969 (B.T) . 5.6 m) individual burrows. 220 they occur in high density. There is no indication of colonialism in areas of high density. They are presumed to be scavengers. In open localities. 1898 COLOR: Carapace dull brown and with conspicuous pits.

sex and the color of substratum of the habitat. HABITS: Construct simple to complex burrows in the soft substratum of the habitat. Garth. Ocypode gaudichaudii H. antennae small and rudimentary. they are confirmed predators and/or scavengers. occidentalis.T) .T) . Rathbun. 1950 (D. 1941b (B) . COMMON NAMES: Cart-driver. 4th pair of legs shorter and thinner than others. Rathbun. eye stalk often prolonged as a style. 1862 COLOR: Upper surface of body and legs generally darkish gray with white marbling. REFERENCES: Alcock. Occasionally occurs sympatrically with 0.T). gaudichaudii: carapace length of 17 mm or more and with well developed ocular styles. 1941b (B). Individual color associated with size. and eastern Pacific (Turtle Bay. All are diurnal and most active following high tide and before flooding midtide. Milne Edwards & Lucas. coral red to dark brown. These two species are distinguishable by: 1) 0. Chace and Hobbs.T) . El Salvador. 7. Crane. 6. HABITS: Burrows highly variable. epistome small. sand-mud areas adjacent to mangrove swamps. also occurs along the shores of lagoons. In several areas species occur sympatrically but generally are distinguishable on the basis of feeding habits or the presence-absence-degree of development of the style over the eye. 2) 0. 1900 (D. Galapagos Islands. Baja California to Chile). 1798 9 CHARACTERS:Carapace deep.T). DISTRIBUTION: Pacific coasts of America. HABITAT: Sandy beaches with tidal surge.1972 BURROWINGLAND CRABSOF THE WORLD Family OCYPODIDAE Genus Ocypode Fabricius. 19 18 (D. Gulf of Fonseca. tips of walking legs and underside of body . 1918 (D. oblique. actively manipulate the substratum feeding on microscopic organic matter (similar to several species of Uca). and 3) straight for 15-20 cm then extending downward in a gradual spiral. Carretero (Peru). coasts of Africa. REFERENCES: Bott. chelipeds shorter than legs and subequal. Crane. somewhat broader than long but generally squarish.T) . Indo-Pacific. Crane ( 194 1) indicates three types: 1) shallow. to Chile. Manus of chelipeds. The second type is most common in the center of the range of distribution. 2) straight with a right angle at 1-3 m depth. Garth. 1843 COLOR: Highly variable. DISTRIBUTION: Tropical and subtropical coast of American Atlantic (Rhode Island to Brazil). simple. Ocypode occidentalis Stimpson. 1969 (G). Red Sea. occasionally on exposed sandy areas when the surge is not high. occidentalis: no ocular style. 1955 (D. rubble flats. 1960 (D). orbits large and divided into two chambers. 1948 (D. HABITAT: Common on sandy beaches of protected bays. Tweedie. Mediterranean Sea.

Baja California to Ancon. Distance from sea generally associated with distribution of same and associated vegetation. 1787) COLOR: Upper surface white with small black spots or generally a pale yellow or a grayish white or a speckled brown. Completely nocturnal in habits except when very young (see remarks in discussion of 0. 1960 (D) . while pale yellow at Punta Cahuita. Brazil. sandy-silt areas adjacent to rivers where water flow is fairly rapid. Garth. Throughout most of Central America. e. Burrows similar to common mode for 0. Crane.45 m. 1969 (G) . due to color. 1918 (D. Feeding begins shortly after onset of ebb until about mid-flood tide. and 2) U-shaped. 1955 (D.10 CONTRIBUTIONSIN SCIENCE No. 1955 (D. dark brown at Tortuguero. Colors tend to vary with substratum except where sand is volcanic (dark black). the U-shaped are most common with the bottom of the U about . Costa Rica.75 m. Chace and Hobbs. sand is then transported in either the right or left cheliped to the surface and then dumped. .T). 8. e. Rathbun.) REFERENCES: Bott. Activity greatest during ebbing of tide and at slack water. HABITAT: Sandy surge-beaten beaches. HABITS: Burrows deep. gaudichaudii). Burrow construction is initiated by a general scratching using the chelipeds followed by removal of sand for the entrance using the dactyli of the walking legs. The general color pattern is apparently associated with the color of the substratum of the habitat.g. Playas de1 Coca. 1941b (B) . Chace and Hobbs ( 1969) give a detailed account of the two color phases found on Dominica.. occidentalis in the Pacific. HABITS: Distinguishable from 0.45-. Chace . ghost crab. including most of the islands in the greater and lesser Antilles.T) . They are scavengers and their rapid movements when disturbed contribute to their common name. Ocypode quadrata (Fabricius. Costa Rica. Costa Rica. COMMON NAME: Ghost crab.. Once the burrow is sufficiently deep. Chace and Hobbs ( 1969) note two types of burrows: 1) vertical or nearly so. DISTRIBUTION: Atlantic coasts of America. These crabs are. HABITAT: Sandy beach areas from upper tidal level to well beyond supralittoral area. Peru. gaudichaudii by absence of ocular styles. but this name is more commonly applied to members of the unrelated genus Emerita. Many show a degree of iridescence along the outer areas of the carapace. COMMON NAME: Ghost crab. or it may be tossed out of the burrow using either cheliped. Rhode Island to Estado do Santa Catharina. (In the past also called a sand crab. 220 cream white. REFERENCES: Bott.T).g. These crabs are scavengers and are probably a geminate species of 0. DISTRIBUTION: Turtle Bay. conspicuous when found on light colored or dry sand. gaudichaudii.

epistome short and distinct. COMMON NAMES: Ghost crab. 1954 (G) . . They have also been reported along the margins of lagoons. 11 1933 (D.T) . somewhat broader than long and with antero-lateral area pronounced and/or projected. Kepiting Mata Panjang (Longeyed crab) (Christmas Island. DISTRIBUTION: American Atlantic (Boston to Uruguay). Tweedie ( 19. including many islands (Mauritius. Ocypode ceratophthalma (Pallas. Chile). 1940 (D) . 1950 (D. Maldive and Laccadive Islands. Millard and Harrison. BURROWINGLAND CRABSOF THE WORLD 1948 (T).T). Note: This has until recently been cited by many authors as 0. Day et al. 1902 (B) .T). Japan. Guam and Hawaiian Islands). Gordon. 1969). Caroline and Marshall Islands.g. HABITS: Burrows are common from the high tide mark to 6-10 m above.D) . including Maury Islands. Sakai. Borradaile. ambulatory legs longer than small cheliped of male and both chelipeds of female. DISTRIBUTION: East coast of Africa (Port St. 1900 (D. 1952 (G) . George and Knott.T). chelipeds of males extraordinarily unequal and large while in female equal and small.. John). In those sites affected by tidal flux the crabs emerge from their burrows during the ebbing of the tide to feed. Tuamotu Islands.. Tesch. Rathbun. and with a splash of yellow on the chelae. Holthuis.T). 1950 (D. Genus Uca Leach. last pair of ambulatory legs shorter than rest. Tweedie. 1880 (D) . but sometimes olive. 1953 (D).. Burrows are generally 37-50 cm deep. REFERENCES: Alcock.T) . 1772) COLOR: Generally the dorsal appearance is from sage green and yellow to grayish white. 1963 (D. and eastern Pacific (British Columbia to Valparaiso. e. 1954). 18 14 CHARACTERS: Carapace deep. Red Sea. 1916 (G). The color is highly variable. Pearse. Well defined ocular styles are present in adults of this species. IndoPacific North to Kii Peninsula. 1802 (see Chace and Hobbs.T). Mediterranean. antennae large and pronounced. west coast of Africa (Portugal to Angola). in the areas above the tidal influence they feed on small organisms. They are scavengers feeding on debris deposited by the tidal exchange. Dakin et al. 1934 (D). Indian Ocean). and associated with the character of the substratum (see Green ( 1964) for discussion of color changes in Hawaiian forms). 9.1972 and Holthuis. albicans Latreille. less frequently they occur in areas beyond the edge of the sea. 1964 (B). Green. HABITAT: Sandy beaches.50) gives the color for those found on the CocosKeeling Islands as uniform gray. 1934 (D). Wrard. 1918 (D. 1954 (B. Barnard. 1918 (D. orbits deep and oblique. Miers. east coast of Africa to Indo-Pacific. crickets (Gressitt.

moderately deep burrows showing considerable variation with age and location. salt marshes. however. 1859) COLOR: No record..T) .T). Salmon. HABITAT: Mud-sandy areas in and adjacent to estuarine situations. Crane. 1914a (B). 1921 (D). Salico&a). b (B). some of these events are correlated with sunset and sunrise as well (see Salmon. 1958 (B). Bott. Salmon. Hagen. Uca subcylindrica (Stimpson. 1965 (B). 1943b. Feeding.g. and 1957) gives a thorough account of display. 1957 (B). low tide muddy areas. DISTRIBUTION: Boston Harbor. This species is restricted by substratum preference. Avicennia. to Brownsville. Most species show considerable social behavior. 1801) COLOR: Carapace cream white. large cheliped of male buff with apricot at base of movable finger. Salmon and Stout. In a typical situation the sand content may increase down a bank from 10 per cent at the top to 60-70 per cent at the low tide level. Dembowski. generally ceasing when the tidal level reaches the burrow entrances. Crane ( 1941 a. 1947 (B) . COMMON NAME: Sand fiddler crab. Many species show a correlation of burrow site with tidal flux whereas others occur widely throughout tidal zone even when burrows may become quite dry. REFERENCES: Barnard. HABITS: They construct well defined. Massachusetts. 1943a (D). Crane. Schmitt. See Pearse ( 1914a) and Dembowski (1925) for details of burrow construction. mangroves. Texas. 11. or pickle weed. There is considerable taxonomic discord associated with this genus due to voids in collections from certain areas and the high degree of variability in color. 220 HABITAT: Associated with a wide variety of mud-sand habitats. burrow repairconstruction and social behavior occur during the low tide period. See Teal ( 1958) for an account of feeding habits as related to sand content. See Salmon (1965) for an additional account of courtship behavior. REFERENCES: Burkenroad. Uca pugilator (Bose. etc. b (B). breeding and relationships of a number of sympatric species of the genus. 1968 (D). b (B). 1954 (D. 10. Area may be exposed or associated with dense vegetation (e. 1943b (B) .g. 1962 (B). DISTRIBUTION: Corpus Christi. clay tidal flats. 1941a (G). chelipeds of small males and females white with a grayish cast. 1965 (B). with the crabs feeding at the lower levels and living near the top. Texas to northern Mexico. mangroves. Teal. 1925 (B). Pearse.12 CONTRIBUTIONSIN SCIENCE No. HABITAT: Sandy areas where sand content is generally more than 40 per cent. . e. HABITS: Most construct well-defined burrows where substratum is moist enough to maintain burrow configuration.. 1965). 1950 (D.

Note: Until recently this genus was placed among the members of the family Gecarcinidae but Chace and Hobbs ( 1969) placed it among the members of the family Ocypodidae. relatively straight. COMMON NAME: Puffed fiddler crab. 1921 (D. Burrows seldom deeper than . Calling crab. REFERENCES: Bright. REFERENCES: Altevogt. e. Uca tangeri (Eydoux. Rathbun. epistome small but prominent. HABITS: Burrows which extend downward for about 30 cm or so are located from the mean tide to highest high tide level. 12. generally shallow. DISTRIBUTION: Portugal. Rathbun. 1897 CHARACTERS:Interorbital distance a little more than one-half the greatest width of the carapace. As the tide recedes the plug is removed and the crab emerges to feed. orbits deep but not much larger than the eyes. Barnard. Hediger. HABITAT: Common inhabitants of muddy shores and mangrove swamps where there is a moderate degree of tidal flux. 1959 (B) . 1961 (D). 1966 (G) . COMMON NAMES: Fiddler crab.T) .T).T). but they often intersect. Juveniles occur in dense numbers per unit area within an interchange of intersecting burrows or in burrows of less density per unit area but where the burrow is directly adjacent and/or attached to those of adults. HABITS: Burrows constructed below the highest high tide level most commonly at midtide level so that burrows covered daily by tidal surge. north and west coasts of Africa to Angola.T) . and constructed in loose sandy soil with high moisture content or in muddy areas. In colonial situations. 1835) COLOR: Carapace reddish brown to dirty yellow. 1962 (B).g. HABITAT: Sandy areas adjacent to brackish water. Habits vary from solitary to colonial. almost always filled with ground water to the level of the burrow mouth. etc. legs stout.75 m. . and is recorded as such by some authors. 19 18 (D. 1934 (B). Genus Ucides Rathbun. Burrows not uniform. the burrows are still simple. slightly twisted.5-10 cm of excavated substratum by the time the burrow is covered by the tide. male large cheliped reddish brown to pale blue. female chelae dirty cream with some pinkish areas. 1962 (B) . 1933 (B). antennules oblique. 19 18 (D. Hagen. Little other ecological data recorded for this species. not common on open coasts. Note: This was described by Eydoux in 1835 as Gelasimus tangeri. See Hagen ( 1961) and Hediger ( 1934) for additional details. The top of the burrow is generally plugged with 7. DISTRIBUTION: East and west coasts of the Americas.1972 BURROWINGLAND CRABSOF THE WORLD 13 HABITS: Burrows are small. salt marsh flats.. 1950 (D. salt marshes. and frequently with multiple entrances. REFERENCE: Rathbun.

Young with a tendency to have a dark gray area along the median anterior margin of the carapace. Juveniles are most often found in small pockets connecting to the burrows occupied by the adults. Rathbun. . HABITAT: Mud of mangrove areas. 14. COMMON NAMES: Wide red land crab. occidentalis in the Pacific. mouths of rivers and brackish water marshes. 1933 (D. in mangroves along mouths of rivers and brackish water marshes adjacent to the sea. 1966 (G) . Burrows are shallow and often with more than one entrance. Burrows do not generally extend more than 50 cm below the surface. Southern Florida (Biscayne Bay) to Santos.g. however. e. Burrows are most common along the mud-water margin of mangroves. 1960 (D). HABITS: This species maintains burrows in areas which are generally covered by high tide at least once per month. dactyli of chelipeds reddish white. Kaburi (Cuba) . cordatus which occurs in the Atlantic. etc. older forms tend to become rust red due to staining from the mud in the burrow. Manning and Provenzano. mostly straight and relatively shallow. HABITS: This crab constructs burrows in very soft mud in areas where there is an absence of low ground vegetation (shrubs). Cangrejo amarillo (Peru). walking legs red-violet. Baja California to Rio Tumbes. Brazil. Molt condition not predictable from color change. DISTRIBUTION: Pacific coast of America (Espiritu Santo Island. Color is variab!e throughout the range of distribution primarily associated with the nature of the habitat substratum. Sulicornia. 1961 (D). Typically.T) . Burrows are wide. Ucides cordutus (Linnaeus. HABITAT: Areas. Peru). de Oliveira. COMMON NAMES: Pagurus.. there is a small side chamber or tunnel paralleling the surface just inside the mouth of the burrow. 1955 (D. Bright. Young and adults construct their burrows in close proximity. 1897) COLOR: Carapace reddish gray with orange-red on the lateral margin. At both ends of the range there is some indication that burrows are shallower and often nothing more than depressions. DISTRIBUTION: Atlantic coasts of America. REFERENCES: Bott. underside brownish white. frequently flooded by tidal surge.T). 1918 (D. may be associated with salt marsh vegetation. 1763) No. This is a geminate species of U. 220 COLOR: Carapace pale yellow with cervical groove and urogastric lobe rusty brown. tips of chelae cream. 1946 (B). 1969 (G). Chace and Hobbs. This is a twin species of U.14 CONTRIBUTIONSIN SCIENCE 13. Primary sites are mangroves with burrows concentrated along the upper edge of distribution of the red mangrove (Rhizophora mangle). Suedu. including the West Indies. The last three ambulatory legs and most of the chelipeds dark red. Garth.T). Ucides occidentulis (Ortmann. Uca (Brazil).


BURROWINGLAND CRABSOF THE WORLD REFERENCES: Bright, 1966 (G) ; Garth, 1960 (D) ; Rathbun,

15 19 18

(D,T). Family GECARCINIDAE Genus Gecarcoidea Milne Edwards, 1837 CHARACTERS: Fronto-orbital border less than half the greatest breadth of carapace; orbits deep; antennae very small and excluded from the orbit; epistome sunken and quite hairy; chelipeds equal or nearly so in both sexes; legs stout. DISTRIBUTION : Indo-Pacific Islands. HABITAT: Moist soil or muddy areas in the jungle areas adjacent to the sea. HABITS : Burrows shallow and not well developed. REFERENCES: Calman, 1911 (B) ; Gibson-Hill, 1947 (B) ; Keilin, 1921 (D); Rathbun, 1918 (D,T); Tweedie, 1947 (D,T); Webb, 1922 (B). 15. Gecarcoidea humei (Wood-Mason, 1873)

COLOR: Dorsal surface a relatively uniform red-violet with some indication of red near the base of the chelae; claws white-brown with reddish violet tinge; scars on carapace yellow or yellowish white. DISTRIBUTION : Indo-Pacific Islands (Nicobars, Andamans, New Britain, Celebes, Christmas (Indian Ocean), Philippines, Loyalties, Formosa, New Guinea, Pulu Weh, and Talauts. HABITAT: Moist areas along the coasts of islands, typically within the jungle. HABITS: Burrows not well developed; generally shallow 15-60 cm in length; mostly parallel with the surface; burrows principally for retreat. Found only where there is a constant source of water to keep soil moist. Andrews (1900) gives a good account of selected aspects of the life history of this species. Gibson-Hill ( 1947) reports extensive migrations to the sea during spawning together with an account of the aspects of development of the young. COMMON NAME: None recorded. REFERENCES: Andrews, 1900 (B); Calman, 1909 (B); A. Milne Edwards, 1879 (T) ; H. Milne Edwards, 1834 (T) , 1837 (T) ; Ortmann, 1894 (D,T) ; Pocock, 1888 (D) ; Rathbun, 19 18 (D,T) ; Sakai, 1940 (D) ; Tweedie, 1947 (D,T), 1954 (B); Wood-Mason, 1873 (T). Note: This is frequently listed as Gecarcoidea Zalandii H. Milne Edwards, 1837. The genus was erected by H. Milne Edwards (1837) for G. Zulandii which was erroneously recorded from Brazil, and then renamed by the same author in 1853 as the genus Pelocarcinus. Wood-Mason ( 1873)) de Man ( 1879)) and Pocock ( 1888) all described this under a number of genera and species. Ortmann ( 1890) reduced all these species to synonymy with Gecarcoidea Zalandii except Pocock’ natalis. Tweedie (1947) gives s a good review of this taxonomic snarl and concludes that G. Zalandii must



No. 220

be regarded as indeterminable on the basis of an incorrect type locality. We use the name humei on the advice of Wood-Mason ( 1873). Tweedie ( 1947) also supports the position that because these are generally island forms there are several variants which can be recognized as subspecies, e.g., the crab found on Christmas Island (Indian Ocean) is G. humei natalis (Pocock) . Genus Cardisoma Latreille, 1825

CHARACTERS Fronto-orbital distance much more than half the greatest : width of the carapace; orbits deep with eyes filling half of the orbit; antennules folded; epistome short and well defined; legs stout. DISTRIBUTION : Tropical America, Cape Verde Islands, west coast of Africa, Indo-Pacific from Port St. Johns, Africa to Hawaiian Islands. HABITAT: Commonly inhabits muddy shores, mangrove swamps and saline lowland soils near the coast, HABITS: Constructs well defined deep burrows in soft soils where ground water is available during the dry season. Often they plug the burrow mouth with mud during the dry season to keep the lower portions of the burrow moist. Burrow sites are always above the mean high tide level. They return to the sea to spawn and to introduce the pre-zoea to the required sea water environment. All are primarily herbivorous but feed on carrion also. REFERENCES: Barnard 1950 (D,T) ; Behre, 1949 (B) ; Bright, 1966 (G); Gifford, 1963 (G); Herreid and Gifford, 1963 (B); Holthuis, 1959 (D); Rathbun, 1918 (D,T); Tesch, 1918 (D,T).
16. Cardisoma guanhumi



COLOR: Carapace deep vio!et in young, but tends to become bluish gray with age or approach to molt; ambulatory legs deep blue with larger cheliped dirty white, Local variation may be due to the type of soil characteristic of habitat. DISTRIBUTION: Atlantic coasts of America, central east coast of Florida, Louisiana, Texas to Florianopolis, Brazil, including the West Indies. HABITAT: Open fields, margins of mangrove swamps, along margins of rivers, in forests, along roads and under houses. In all known situations the soil is saline. HABITS: All are typically found within a few hundred yards of a brackish or saltwater source. The young are found under debris, e.g., coconut husks, palm fronds, flotsum, etc., directly adjacent to salt water (generally above the highest high tide level) or in the burrows of adults. Adults construct burrows of varying depth and structural complexity, depending upon their age and the location of the burrow with respect to available ground water. Large aggregations of adult crabs are often found in areas where the substratum is soft yet still suitable for burrows, and where there is little ground cover. Although adults occur several hundred meters from the sea, they are most common within about 200 meters of the tidal zone. Inland dwelling




individuals migrate to the sea in great numbers during the breeding season to shed their eggs. This is a twin species of C. CY~SSU~which occurs in the Pacific. COMMON NAMES: Great land crab; White land crab, Juey, Cangrejo; Guanhumi; Mulatto land crab; Guaiamu, guarani, guayamu (Brazil). REFERENCES: Behre, 1949 (B) ; Bott, 1955 (D,T) ; Bright, 1966 (G) ; Chace and Holthuis, 1948 (T) ; Feliciano, 1962 (B) ; Gifford, 1963 (G); Herreid, 1963 (B), 1967 (B) ; Herreid and Gifford, 1963 (B) ; de Oliveira, 1946 (B); Pearse, 1916 (G); Peyton et al., 1964 (D); Rathbun, 1933 (D,T). 17. Cardisoma
crassurn Smith, 1870

COLOR: Carapace deep blue; dactyli of walking legs red; large cheliped pale yellow to dirty white; underside cream-white. DISTRIBUTION: Pacific coasts of America. Todos Santos, Baja California to the Rio Chira, Peru. HABITAT: Open fields, margins of mangrove swamps, along roads and fence-rows, margins of rivers and streams, under houses and in cultivated fields; generally in saline lowland soils near the coast. HABITS: These show habits similar to those of C. guanhumi in areas adjacent to brackish or salt water sources. In contrast to C. guanhumi the young construct separate shallow burrows along river banks and edges of mangroves. Adult burrow construction parallels that of C. guanhumi. During the dry season, adults with burrows in open, exposed areas close the top of the burrow with a plug of mud. Some reports have indicated that closure of the burrow also occurs prior to the onset of ecdysis (shedding). Adult migrations to the sea during the spawning period are common in Mexico, Costa Rica, Panama, and Ecuador. This is a geminate species of C. guanhumi which occurs in the Atlantic. COMMON NAMES: Mouthless crab, Cangrejo sin boca (Peru), Cajo (Mexico). REFERENCES: Bott, 1955 (D,T) ; Bright, 1966 (G) ; Garth, 1948 (D,T), 1960 (D); Murphy, 1944; Pesta, 1931 (D,T); Rathbun, 1918 (D,T). 18. Cardisoma

Herklots, 185 1

COLOR: Young, newly molted individuals with violaceous carapace; tips of chelae and walking legs bright red; with age and approach to molt carapace turns dirty yellow with occasionally slight reddish spots dorsally. DISTRIBUTION: Western coast of Africa from St. Louis to Baia dos Tigres, Angola, Africa and Cape Verde Islands. HABITAT: Moist sandy areas above the mean high tide level; mangroves, mouth of rivers, under houses, in cultivated areas adjacent to permanent sources of brackish or sea water; and inland areas of larger islands. HABITS: Youngest juveniles are in small depressions or newly dug shallow burrows directly adjacent to water; older juveniles found in smaller compartments within the burrows of adults. Adults construct deep burrows,

including Mauritius. Kepiting Balong (Cocos Island). e. mud banks in the immediate neighborhood of fresh-water. Papaka Tupa (Tuamotu Islands). 1902 (B) . Rathbun. Wanson. Andaman Islands. HABITS: Constructs well defined burrows in soft soils where ground water is available during the dry season. 1950 (D. Holthuis. 1960 (B) .T) . Not uncommon in the jungle adjacent to the sea. hirtipes (see Miyake.D).T) . 1939). Hogue and Bright. 19 18 (D. Barnard. Forest and Guinot. often moving considerable distance from their burrows to feed on palmnuts. 1922 (B). . 19 10 (D) . 19. Borradaile. Madagascar. Cheesman. On coral atolls it is common among coconut husks. north eastern Australia and north toward Japanese Mainland (Loo Choo). Tweedie ( 1947) notes that on Christmas Island (Indian Ocean) the carapace is light bluish gray and the chelae are dirty white. under rubble piles and in mixed forest areas adjacent to plantations. REFERENCES: Alcock. Miyake (1939) gives a good account of the distinguishing species characteristics. Cardisoma carnifex (Herbst. 1950 (D. Habits generally parallel other members of the genus.T) . whole of Indo-Pacific.. HABITAT: Common inhabitants of muddy shores. 1939 (D. 1950 (D.T) . There is considerable color variation throughout the Indo-Pacific. chelipeds light purple to dark cream. Stebbing. 1900 (D. DISTRIBUTION: East coast of Africa. 197 1 (B) .T) .g. DISTRIBUTION: Occurs in sympatry with C.18 CONTRIBUTIONS IN SCIENCE No. Cardisoma hirtipes Dana. Both juveniles and adults are nocturnal scavengers. Tesch. Dalziel. mangrove swamps or Kuli and saline lowland soils near the coast. 1920 (D). 1963 (B. dead fish and scraps of vegetation. HABITAT: Moist saline soils. 1794) COLOR: Carapace dark purple.D). Silas and Sankarankutty. 1935 (D). COMMON NAMES: Land crab. Malay Archipelago. 1923 (B) . 196 1 (D). Spawning activities have not been recorded in the literature for this species. Note : Over much of the range this occurs in sympatry with C. 1953 (D). 220 and often these are part of a large colony where the burrows intersect. 20. Miyake. REFERENCES: Barnard. coconuts. Tweedie. 1966 (B. Where soil is dense or crusted they frequently scratch-out a space under a treeroot or rock. All around the Indian Ocean it is most commonly found between the high tide mark and just beyond the extreme highest high tide line. Polynesia and Melanesia. 1851 COLOR: Carapace generally dark violet and chelae bright cinnamon red.T) . COMMON NAME: Edible land crab. Macnae. 1921 (D.T). carnifex from east coast of Africa throughout whole of Indo-Pacific.

hirtipes would probably result in extension of this distribution. antennae very short. 1940 (D). the second pair being longest. 2 1. legs stout. to specific status from synonymy with Cardisoma hirtipes. river mouths and adjacent coastal sandy and saline soil areas. occasionally causing crop damage to melons and pumpkins (Esaki. Garth. In the extreme northern and southern portions of the distribution the burrows are deep 1. A few weeks after the onset of the rainy season in January or February spawning occurs. and further states that examination of series presently considered as C. Villalobos and Cabrera. 1918 (D. Chace and Hobbs. Australasia. Sakai. Finnegan. 1948 (D. Restricted to islands from west coast of Mexico to Colombia. 1853.T) . Perhaps it is the ecological equivalent of the American species of the genus Ucides. DISTRIBUTION: Pacific coasts of America.T). REFERENCES: Alcock. except during rain storms. 1966 (G) . DISTRIBUTION: Tropical America. 193 1 (D) . 1947 (B). Esaki. . Gecarcinus planatus Stimpson. The burrows are only SO-75 cm below ground level. Gibson-Hill. Many utilize debris as a source of protection in lieu of a burrow.1972 BURROWING LAND CRABSOF THE WORLD 19 HABITS: Normally the crab digs burrows in the soft mud directly adjacent to streams. HABITAT: Drier areas above the tidal margins of mangroves. 1940)) but in areas associated with human habitation they are carrion feeders as well. They are primarily plant feeders. 1964 (B) . Considering our preliminary studies on related species it seems likely that this species can abide in areas where there is more ground water than is characteristic for C. 1939 (D.T) . hirtipes. Genus Gecarcinus Leach. Milne Edwards. He gives the distribution of C. 1860 COLOR: Body and legs generally an orange-red. tips of chelae cream with small flecks of brown. frontalis as Loyalty Islands. Note: Tweedie ( 1950) restored Cardisoma frontalis H. northern Daitozima. tips of walking legs often dark red. 1969 (G) . West and South Africa. Tweedie. epistome linear. Miyake.T). Bermudas. carnifex. HABITS: Burrows always shallow and devoid of ground water. 1947 (D. However. COMMON NAME: Land crab. Copulation occurs at the edge of the sea just prior to the time the female sheds the previous batch of eggs.T). 1900 (D. 18 14 CHARACTERS:Fronto-orbital distance half or less than half of the greatest width of the carapace. until there is an extensive revision of the genus with clarification of the number of island endemics we will herein consider these still to be C.2 m and often with mouth plugged during the dry season. 1940 (D) . Tesch. REFERENCES: Bright. and Cocos-Keeling Islands. Japan. Ascension Island. orbits deep with eyes nearly filling the orbits. Gordon. 1934 (D).

During the rainy season they are reported to move in large numbers to the sea to breed. 1918 (D. e. 1969 (G) .. southern Florida. REFERENCES: Chace and Hobbs. often associated with beach strand vegetation HABITS: Adults and older juveniles scratch out shallow burrows under rocks. Big red crab. they are common along mountain slopes and cliffs adjacent to the beach and to heights of 500 meters and as far as a thousand meters from the shore. however.T). Isla Providencia. which are quite frequently under a tree or the edge of a large rock. Burrow serves primarily as a hiding place. under roots and in soft soils above highest high tide mark on slopes up to 120 m. 23. Along the shore edge. Gecarcinus quadratus Saussure. Curacao. After metamorphosis. 1918 (D. Cangrejo rojo (Panama).T). young birds. and Cayman Islands. greater and lesser Antilles. Gecarcinus ruricola (Linnaeus. very shortly after the second or third molt they move to areas well above the highest high tide. the young are found in large numbers just above the high tide level. Colombia. COMMON NAMES: Island crab. Newly metamorphosed juveniles hide along the shore under debris. They are more secretive than most of the gecarcinids. planatus in the Pacific. large chelipeds with light purple tinge. Burrows have no standing water. They are nocturnal feeders.20 CONTRIBUTIONS IN SCIENCE No. Young hide in natural crevices and small spaces providing natural protection.g. ambling over almost anything in their path. sleeping scientists.g. COMMON NAMES: Black crab. Restricted to islands: Bahamas. REFERENCES: Garth. HABITAT: Low and marshy areas not far from the sea. 1758) COLOR: Body and legs generally black with purplish tinge. 1960 (D). e. 1948 (T). On larger islands. Mountain crab. lower slopes of island mountains up to 500 meters. Red tourlourou. 220 HABITAT: Rocky areas. etc. 1948 (D. last two joints of legs red. HABITS: They hollow out obliquely inclined shallow burrows. older individuals or those undergoing late preecdysial changes are overall much lighter in color. small light yellowish spot on the posterior margin of the carapace. Blue land-crab. and commonly move considerable distance from their burrows. Rathbun. Rathbun.T). ruricola occurring in the Atlantic.. often gregariously. Chace and Hoithuis. 1853 COLOR: Carapace brownish red with two white spots in the cardiac region. they are often found sympatrically with Gecarcinus lateralis. food lockers. This is a geminate species of G. intestinal region orange-red. This is a geminate species of G. roots or debris. . DISTRIBUTION: Atlantic coasts of America. 22. Bold when on feeding excursions. abdomen light yellow with violet hue. red and yellow patch below the orbit of the eye.

. 1955 (D. coconut husks and fronds. Zpomoea. 1965 (B) . REFERENCES: Bright. Ray. and low growing vines. coconuts.. Pattern of dark red carapace highly variable throughout range of distribution and in distinct (isolated) populations. etc.T) . underside sooty white.. Zpomoea (family Convolvulaceae) . Mexico to La Libertad.. There is also a tendency for the depth of the burrow to be correlated with the length of the dry season. REFERENCES: Bliss. Also occurs on islands in the West Indies. Burrow construction is correlated with the length of the dry season. Crabs at both extremes of the distribution tend to construct simple burrows. (Atlantic side of Colombia).T). i. COMMON NAMES : Black land-crab. 1948 (T). Cabrera. HABITAT: Above the highest tide zones of sandy beaches in moist forest and mangrove areas where there is low growing vegetation or debris. Pesta.T).T) . e. 1948 (D. 1918 (D. Burrow construction is as for G. 193 1 (D. This is a geminate species of G. Ecuador. Not known to occur on permanently isolated islands. 1916 (G). where there is no standing water but a good bit of interstitial soil moisture.5 . burrows are not common. HABITS: All occur in drier areas directly adjacent to mangroves or the ocean. Chace and Holthuis.g. Yucatan. 24. Burrows are deeper on the extremes of the range of distribution.. associated with a variety of beach strand vegetation. Common land-crab. South Padre Island. Texas. Florida Keys. underside cream-white.g. Bahamas. to Macuto. 1964 (B) . Acapulco.g. Non-burrowers tend to occupy small depressions under vegetation debris.1972 BURROWINGLAND CRABSOF THE WORLD 21 merus of maxilliped light yellow. e. Pearse. Bott. Chace and Hobbs 1969 (G) . Rathbun. later-ah in the Atlantic.T) . 1966 (G) . e. HABITS: All occur in nearly dry areas. ARTHROPOD INHABITANTS OF LAND CRAB BURROWS The following list represents an attempt to cite all published records of arthropods found in land crab burrows. 1933 (D. Because the host was not identi- . 7-50 cm deep. DISTRIBUTION: Atlantic coasts of America. 1966 (G) .15 m above the supratidal area where there is a dense covering of debris. They are most common along the uppermost areas of sandy beaches from 1. Bright. while at midrange. Gecarcinus lateralis (Freminville. COMMON NAMES: Red land crab. quadratus. e. Finnegan. dactyli sooty gray. Turbo. HABITAT: Along the upper dry zone of sandy beaches and adjacent low hills. 6 to 9 m above highest high tide level. Venezuela. or low growing beach strand vegetation. Garth.g.e. DISTRIBUTION : Primarily Pacific coasts of America. Whitespot crab. 193 1 (D) . 1835) COLOR: Carapace dark red with small white spots just posterior to the eyes and a pair of white spots in the cardiac region. 1967 (D). houses. chelipeds reddish gray.

To this list could be added several more from unpublished works known to us and no doubt others from other studies now under way.. lobster. Accounts of dubious validity are also included for completeness and to establish the need for verification. Our bibliographic research would indicate that. The list has been read by various specialists and it is hoped that errors of identification. 1928 : tropical America. etc. Dyar. Presently 140 species of mosquitoes are recorded as either resting as adults or breeding in crabholes. with a few notable exceptions. ( 1965: 37-38). Mattingly. 1959: Indomalayan Region). Most of these transient crabhole breeders are salt water adapted or tolerant species which are general ground pool breeders along the coast. 1966: Papuan Region.22 CONTRIBUTIONS IN SCIENCE No. Barraud. . Crabholes are easily sampled using the crabhole pump and collection methods described by Belkin et al. crab. references to all are included. 1952: Ethiopian Region. We feel that much more attention should be paid to crabholes in general mosquito surveys than has been customary in the past (see remarks of Peyton. 1970: 2-3). 1958. INSECTS Order DIPTERA Family CULICIDAE We found it impossible to scour the voluminous literature on mosquitoes for all records of species utilizing crabholes. the determinations of the real crab owner of the burrow from which the associates were collected. Hopkins. seem to regard the crabhole as an aberrant breeding site being utilized by only a handful of specially adapted species. 220 fied in all cases and there is frequent confusion in usage of the terms land crab. whereas those taxa specifically adapted to crabholes are indeed few. Some explanation is in order regarding certain assumptions made and conventions used in compiling the list. Steffan. comprehensive (though not always current) reviews including ecological data are available (Belkin. 2) The type of association and degree of dependence (see Table I and discussion of the Crabhole Community above) of the species on the crabhole were ascertained or inferred from all available information on the biology of the species. Dubious decisions and the criteria for allocation to a category are explained where relevant. erroneous records and synonymies have been largely detected. are assumed to be correct.. for most regions.e. Fortunately. 1) Identifications and associations with the host crab. Surprisingly no culicidologist has attempted previously an exclusive investigation of this habitat. the number of transient species is much larger than previously suspected. Delfinado. Most authors. 1934: India. 1966: Philippines. 1962: South Pacific. i. crayfish. mud lobster.

At least a few crabhole mosquitoes are of known public health importance. as have various other mosquitoes which develop in crabholes.. 1970). CRAB HOST: Cardisoma armatum. 1961:19). including Deinocerites. but by no means universal. Lab. Templis and Galindo. Grayson. 1956). the adaptive significance of which is totally obscure. 4. among larvae which develop in waters with high salt content. From this it seems probable that the specific crabhole mosquito fauna is of mixed origin derived from both of these more primitive categories through convergent adaptations (van den Assem. merus) for both of which there are several well authenticated records of breeding in crabholes (see list). etc. The nature of these adaptations is virtually unstudied.. and in the south Pacific. Stubby or vestigial anal papillae on the larvae. especially in mangroves. 1967). pseudes in Panama (Galindo. These species are Aedes aegypti and Anopheles gumbiae (? melas. interior and coastal. coral rock pools. 19 10) DISTRIBUTION : Tropical Africa. and artificial containers) also are found in the crabhole. tidal pools.. puddles. Aedes pembaensis. A considerable number of container breeding species (tree holes. The former species has also been found to harbor several kinds of viruses of unknown but possible pathogenicity (Heisch et al.1972 BURROWINGLAND CRABSOF THE WORLD 23 Their usual normal habitats consist of salt marshes. This condition is common. 1970: 175. 3. while not specific or even semispecific members of the crabhole community. Two primary vectors of serious diseases in Africa. mangrove pot holes. 1967) and oviposition directly on the host crab in A&es pembaensis (Goiny et al. 1957). Vectors of filariasis on the east African coast. are semispecific members of the crabhole community. Trypanosome organisms have also been recently isolated from wild adults of this species in Panama (Gorgas Mem. . Short head hair l-C. Aedes polynesiensis. 2. While Deinocerites do not appear to be strongly anthropophilic they may act as important agents in maintaining virus reservoirs in coastal animal populations (silent cycles) and which may enter the human population via other vectors. The eastern Equine. Genus Aedes Subgenus Aedimorphus A. Very specialized and aberrant reproductive behavior such as pupal attendance and lack of swarming in Deinocerites (Provost and Haeger. 1967. Louis encephalitis viruses have all recently been isolated from D. nevertheless may develop in tremendous numbers in this habitat and may even find refuge there during eradication programs designed to treat only the more usual breeding sites. Prolonged developmental period. Venezuelan and St. Some characteristics frequently observed in crabhole species are as follows: 1. abnormalis (Theobald.

domesticus (Theobald. 1935:576-578. Seychelles. 1917: 142 (and as minutus). saline seepage pools on coast. in interior. Note: Identification dubious. D-distribution. TYPE: Transient. IUnless followed by symbols indicating additional significant information (T-taxonomy. REFERENCE: Wanson. Madagascar. Usually breeds in grassy swamps. etc.. Usually breeds in fresh water ground pools. A. REFERENCE: Dalziel. REFERENCE: Wanson. A. TYPE: Transient. centropuncta tus Theobald. Ingram and Macfie. coastal. A. 579. the references cite only records of the occurrence of the species in crabholes. B-biology.24 CONTRIBUTIONS IN SCIENCE No. Probably breeds in ground and rock pools. interior and coastal. G-general). Recorded also from borrow pits and stream pools. REFERENCEI: Wanson. 1910) DISTRIBUTION : Nigeria. CRAB HOST: Cardisoma armatum. 1901) DISTRIBUTION : Tropical Africa. 1903) DISTRIBUTION: Tropical Africa. durbanensis (Theobald. Adults only. Arabia. 1960 :99. British West Africa. 19 13 DISTRIBUTION: Sudan. caliginosus (Graham. pools. CRAB HOST : Sudanonau tes africanus. 220 TYPE: Transient. interior and coastal. borrow pits. A. interior and coastal. A. A. 1935: 576. 1920: 253. etc. CRAB HOST: Cardisoma armatum. albocephalus (Theobald. primarily coastal. Normally breeds in grassy swamps. interior by watercourses (?) . Adults only. fowleri (Charmoy. 1952: 182. CRAB HOST : Not recorded. CRAB HOST: Cardisoma armatum. REFERENCES: Hopkins. CRAB HOST : Cardisoma armatum. TYPE: ? Bionomics insufficiently known. 1908) DISTRIBUTION : Africa. REFERENCE: Hanney. TYPE: Accidental. 1935:576-577. TYPE: Transient. . 1903) DISTRIBUTION : Africa.

? Thailand. Normally breeds only in ground pools. CRAB HOST: Cardisoma armatum. TYPE: Accidental. None of the crab hosts has been identified. 1922 DISTRIBUTION : Andaman Islands. CRAB HOST : Cardisoma armatum. 1935. 190 1) DISTRIBUTION: West Africa. 1935:576. TYPE: Semispecific. REFERENCE: Wanson. REFERENCES: Kumm. irritans (Theobald. 1907) DISTRIBUTION: Tropical Africa. 19 15 DISTRIBUTION: Borneo. Dalziel. Note: May be confused with centropunctatus. Wanson. primarily coastal. Wanson. Kumm. 1920:251-253. Ingram & Macfie. tarsalis (Newstead. 1935. The entire subgenus appears to be adapted to this habitat and comprises a specific or semispecific member of the crabhole community though mangrove pot holes and coastal ground pools may serve as secondary breeding sites.1972 BURROWINGLAND CRABSOF THE WORLD 25 TYPE: Transient. Subgenus Cancraedes GENERAL REFERENCE: Mattingly. 195 1: 119.120 (B) . punctothoracis (Theobald. Dalziel. *A. cancricomes Edwards. REFERENCES: Bruce-Chwatt and Fitz-John. Philippines. Kumm. 1920 : 251. All species have a coastal distribution or are found on small islands. Dunn. TYPE: Transient. All species for which the immatures are known (*) are found breeding primarily in crabholes from which all have been taken as adults. Malaya. 195 1: 119. 1920:253. A. Usually breeds in rock and ground pools. Dunn. A. A. 579 (as nigeriensis) . curtipes Edwards. . A. Strongly anthropophilic. interior and coastal. Wanson. 1917:135. Adults only. 1935 :576-577. CRAB HOST: Cardisoma armatum. TYPE: Semispecific. 1931:65. Wanson. CRAB HOST : Cardisoma armatum. Also breeds in ground pools. 1931:65. 193 1: 65 (as sudanensis) . 1901) DISTRIBUTION: West and Central Africa. REFERENCES: Bruce-Chwatt and Fitz-John. 1916:7 (as sudanensis). 1935:576-577. 1928:249. primarily coastal. coastal. Macfie and Ingram. Breeds also in small brackish pools along the coast. 1928:249. A. REFERENCES: Dalziel. nigricephalus (Theobald. 1958. Usually breeds in rock pools and grassy ground pools. 19 10) DISTRIBUTION: West tropical Africa.

A. A. baisasi Knight and Hull. longiforceps DISTRIBUTION : Solomons. mamoedjoensis DISTRIBUTION : Celebes. kohkutensis No. A. thurmanae Mattingly. indonesiae Mattingly. becki Belkin. *A. 1958 1958 1958 1958 DISTRIBUTION : Thailand. simplex (Theobald. east Gulf of Siam. 1958 DISTRIBUTION : Celebes. 1968 1903) DISTRIBUTION : Taiwan. 1959. ? Philippines. DISTRIBUTION : Malaya. A. A. None of the crab hosts has been identified. masculinus Mattingly. Subgenus Geoskusea GENERAL REFERENCES: Mattingly. 19 16) DISTRIBUTION : Australia. DISTRIBUTION : Philippines. daggyi Stone and Bohart. New Guinea. palawanicus Mattingly. Thus all can probably be classified as specific or semispecific members of the crabhole community. the adults of all species have been taken from this habitat. Solomons. 220 Mattingly. fimbripes Edwards. 1924 Brug. Sumatra. *A. DISTRIBUTION : Ceylon. A. A. *A. kabaenensis DISTRIBUTION : Celebes. Mattingly. A. 1929 DISTRIBUTION : Bismark Archipelago. 1962 DISTRIBUTION : Solomons. 1939 Edwards. dnliensis (Taylor. 1958 DISTRIBUTION : Java. 195 1 DISTRIBUTION : Philippines. . A. *A. 1962:332-339. A. 1944 DISTRIBUTION : New Hebrides. As with the preceding. crabholes in coastal areas are the primary breeding places of all species in this subgenus for which the immatures are known (* ) . Belkin. penghuensis Lien.26 CONTRIBUTIONS IN SCIENCE *A.

1927) DISTRIBUTION: Australia. 1901 DISTRIBUTION : Jamaica. 1924 DISTRIBUTION : Moluccas. tropical Africa. REFERENCES: Amos. A. thalmus musgravei) A. CRAB HOST: Cardisoma armatum. REFERENCE: Wanson. 1969 : 35. Larva predaceous. Belkin. walkeri and broken Theobald. 1907) DISTRIBUTION: Jamaica. 579. REFERENCE: Berlin. Larva predaceous. interior and coastal. interior and coastal. China. suvae Stone and Bohart. New Guinea. tonsus Edwards. Normally breeds in bromeliads. . Subgenus Mucidus A. Burma. coastal. inaequalis (Grabham. A.” TYPE: Specific. TYPE: Transient. 1969 :48. Usually breeds in transient ground pools and marshes. TYPE: Transient. “crab pot hole. REFERENCE: Steffan. interior and coastal. 1965). A. 1935:576-577.1972 Note: BURROWINGLAND CRABSOF THE WORLD 27 Adults have been observed feeding on mud skippers (Periophresting on mangrove roots in the Solomon Islands (Sloof and Marks. coastal. CRAB HOST: Not recorded. perryi Belkin. Subgenus Howardina A. Most commonly breeds in treeholes bamboo. 1966:206. 1944: 32. 1962 DISTRIBUTION : Solomons. No other recorded breeding site. scatophagoides (Theobald. CRAB HOST: Not recorded. Ceylon. CRAB HOST: Not recorded. 1944 DISTRIBUTION : Fiji. aurantius chrysogaster (Taylor. “crab and lobster holes. CRAB HOST: Not recorded. TYPE: Transient. Usually breeds in various types of ground pools in coastal areas. Subgenus Levua A.” TYPE : Transient. 1962: 400 (G) . REFERENCE: Berlin. 190 1) DISTRIBUTION: India.

Forattini. 1958: 177-178. Adults only. perventor Subgenus Ochlerotatus Cerqueira and Costa. tropical Africa. Montchadsky and Garcia. A. REFERENCE: Mattingly. prefers saline water? REFERENCE: Delfinado. 1968: 33. Strongly anthropophilic. Bionomics poorly known. tide pools. A. 1967 :20. A. REFERENCES: Belkin et al. 19 12: 19 (as Culex taeniorhynchus) (B) . fronds also . TYPE: Transient.. Subgenus Neomelanoconion A. 182 1) DISTRIBUTION: American coasts and interior saline areas. 220 DISTRIBUTION : Southeast Asia. and inland saline sinks. 1970:49 (G). REFERENCES: Delfinado. Lutz. CRAB HOST: Cardisoma guanhumi. 1905) DISTRIBUTION: Oriental Region. 1914 DISTRIBUTION : Philippines. 1946 DISTRIBUTION: Brazil. coastal. interior and coastal. 1958 DISTRIBUTION : Malaya. Adults only. panayensis Ludlow. 1966: 214. 1966:42. TYPE: Transient. Australia. de Oliveira. Subgenus Paraedes A. TYPE: Transient. Steffan. CRAB HOST: Not recorded. 1958:37 (B). etc. bonnae Mattingly. Zineatopennis (Ludlow. Usually breeds in marine littoral ground pools. 1958: 175-177 (B). Forattini et al. Adults only. Breeds usually in vegetated ground pools... Usually breeds in puddles and hoof prints near the coast. CRAB HOST: Cardisoma guanhumi. coastal. Only breeding records from crabholes. taeniorhynchus (Wiedemann. Moluccas. CRAB HOST: Not recorded. 1946:297. TYPE: Probably transient. REFERENCES: Forattini. REFERENCE: Wanson.28 CONTRIBUTIONS IN SCIENCE Subgenus Neomacleaya A. TYPE: Transient. usually found in coastal salt marshes. CRAB HOST : Not recorded. 1935: 576. TYPE: Transient. CRAB HOST: Not recorded. 1958 : 34. Salt water breeder. interior and coastal. New Guinea. Usually breeds in ground pools (palm recorded). dux Dyar and Shannon. 1925 No. coastal.

TYPE: Transient. albomarginatus (Newstead. Mattingly. 1952 :224.. The females deposit their eggs on the legs and body of the host. 1958 :40. Seychelles. Subgenus Skusea A. also commonly utilizes ground pools and swamps and rarely natural and artificial containers. Usually breeds in numerous other marine littoral ground pool habitats. Goiny and Ikata. lunulatus King and Hoogstraal. REFERENCE: King and Hoogstraal. from crayfish hole in shaded rain forest. coastal. Subgenus Rhinoskusea A. after Mattingly. 1907) DISTRIBUTION: Central tropical Africa. north Australian and Papuan Regions. Hogue and Bright. 250 feet elevation. Lumsden. especially mangrove swamp pools. coastal. eulimene. and artificial containers. 1946 DISTRIBUTION : New Guinea. CRAB HOST: Not recorded. TYPE: Semispecific. 1908) DISTRIBUTION : Indomalayan. Leicester. 1955: 170-171 (B).1972 BURROWING LAND CRABSOF THE WORLD 29 Subgenus Pseudarmigeres A. 1958 : 39-40. CRAB HOST: Not recorded. pillaii Mattingly. 190 1 DISTRIBUTION : East Africa. coastal. S. 1957 : 144. 196 1 (B) . 1935:577. interior and coastal. Subgenus Pseudoskusea A. 1946a:97. “crayfish hole. REFERENCES: Brook Worth et al. Madagascar. CRAB HOSTS: Sesarma meinerti. Usually breeds in numerous other marine littoral ground pool habitats. REFERENCE: Reid in litt. pembaensis Theobald. 1958 DISTRIBUTION: Malaya. 197 1) . REFERENCES: Colless. TYPE: ? Single breeding record. coastal. 1957). Females are strongly anthropophilic and vectors of filiariasis in east Africa (Heisch. TYPE: Transient. from crabhole. longirostris (Leicester.” TYPE: ? Single breeding record. (Goiny et al. A. REFERENCE: Wanson. Hopkins. CRAB HOST: Cardisoma arm&urn. Predominantly breeds in crabholes. 1957. CRAB HOST : Not recorded. 1908 :8 (as Ficalbia Zongirostris) .. .

1962) DISTRIBUTION : Cosmopolitan. 1935:576. Usually breeds in containers of various sorts REFERENCE: Belkin. coastal. 2. 195 1 DISTRIBUTION: South Pacific islands. 220 (Linneaus. rockholes and temporary ground pools). polynesiensis Marks. CRAB HOSTS: Cardisoma carni. CRAB HOSTS: Cardisoma armatum. Normally breeds in treeholes (also in cut bamboo. hirtipes. 1920:251-252. CRAB HOST: Not recorded. coastal. 1929. 1935:576 (as argenteus). REFERENCES: Bruce-Chwatt and Fitz-John. CRAB HOST: Not recorded. 190 1) DISTRIBUTION: Tropical Africa. interior and coastal. TYPE: Transient. 1928:249. aegypti No. 195 1: 120. TYPE: Transient. interior and coastal. 1962:468. TYPE: Accidental. pseudoscu telluris (Theobald. interior and coastal. 1935: 576-577. A. Riqueau. 1934:590-593. 1960:5. Sesarma africanum. luteocephalus and (Newstead. 1960:5. REFERENCES: Edwards. Dalziel. CRAB HOST: Cardisoma armatum. pl. Strongly anthropophilic and vector of yellow fever. C. Usually breeds in containers of various sorts. Tamashiro. TYPE: Semispecific. africanus (Theobald. 1908 : 8. coastal. Cheneveau. 251-252. Symes. 1960. Burnett. 8. A. Usually breeds in various artificial container habitats. butleri Theobald. 1920:248. 1962: 470. 1907) DISTRIBUTION: Tropical Africa. Strongly anthropophilic and vector of filariasis and dengue. . Subgenus Verralina A. Leicester. Anthropophilic vector of yellow fever. 8. 1901 DISTRIBUTION: Indomalayan Region. CRAB HOST: Cardisoma armatum. 19 10) DISTRIBUTION : Fiji. 1928 :346 (as umbrosus) . TYPE: Probably semispecific. Wanson. REFERENCE: Wanson. REFERENCES: Belkin. Most commonly known from ground pools in mangroves and nipa palm axils. pl.fex. 1964: 10-l 1 (B). 2. Dunn.30 CONTRIBUTIONS IN SCIENCE Subgenus Stegomyia A. A. Crabholes important breeding sites when other habitats absent as on low coral islands. Wanson. Normally breeds in treeholes. A. TYPE: Transient or accidental. REFERENCES: Dalziel. Symes.

” TYPE: Transient. 1935:576. Usually breeds in various types of ground pools. 19 14) DISTRIBUTION : New Guinea. “crayfish hole. complex. 1961:25. normally breeds in vegetated ground pools and sluggish streams. South America. 1902. 1821 DISTRIBUTION: Tropical America. interior and coastal. Dunn. Usually breeds in a wide variety of artificial and natural ground habitats. TYPE: Accidental. 1917:393-394. Anthropophilic and a vector of malaria. REFERENCES: King and Hoogstraal. CRAB HOST: Not recorded. 1947 DISTRIBUTION: New Guinea. Wiedemann. tigertti Scanlon and Peyton. Dalziel. 1902) or melas (Theobald. 1967 DISTRIBUTION: Thailand. Bruce-Chwatt and Fitz-John. TYPE: Semispecific or specific.. 1964). 1916:7. Solomons. REFERENCE: Scanlon and Peyton. interior. 578. Note: The salt water species (?) merus (Donitz. Subgenus Nyssorhynchus A. 1927 :3 3 8 (as tarsimaculata in part) . Flushed into crabhole by heavy rains. 1963: 10. . Subgenus Cellia A. van den Assem. 19 17: 135 (as costalis) . Fresh water species. CRAB HOST: Not recorded. Cheneveau. REFERENCES: Aders. 1967. albimanus CRAB HOST: Not recorded. REFERENCES: Peters. Sesarma africanurn. primarily coastal? CRAB HOST: Not recorded. 1934:590-593 (as cost&is). Macfie and Ingram. CRAB HOSTS: Cardisoma armatum. 1970:49.r King and Hoogstraal. Ingram and Macfie. Bismark Archipelago. 1947: 125. 1903) may be found ultimately to be those associated with crabholes (see Coluzzi. TYPE: Accidental. breinli (Taylor. 1920:251-253. 1928: 249. Adults only. REFERENCES: Shropshire and Zetek. TYPE: Transient.1972 BURROWING LAND CRABS OF THE WORLD 31 A. primarily coastal. gclmbiae Giles. Species breeds in plant containers and rarely ground pools. Belkin et al. Steffan. Genus A rmigeres A. interior and coastal. parasimi2i. Genus Anopheles Subgenus Anopheles A. 1966:215. 1951: 120. DISTRIBUTION: Africa. Wanson.

Dyar and Knab. TYPE: Transient. C. 1906) No. 1935:576. 220 DISTRIBUTION: Guadeloupe Island. C. Known only from crabholes. TYPE: ? Bionomics poorly known. Multiple collections. REFERENCE: Floch and Abonnenc. 578. all from crabholes. 2.32 CONTRIBUTIONSIN SCIENCE Genus Culex Subgenus A edinus C. corrigani Dyar and Knab. usually breeds in ground pools but also occurs commonly in almost all other habitats. annulirostris Skuse. Lesser Antilles. 1932:97. Forattini. Dyar and Knab. coastal. CRAB HOST: Not recorded. Lutz. Wanson. Ucides cordatus. TYPE : Specific. 1912:19. 1956:85. interior and coastal. REFERENCES: Howard. Rabello and Heredia. annulioris Subgenus Culex prob. 1945: 39. TYPE: Specific or semispecific. Usually breeds in ground pools. 1907 DISTRIBUTION: Panama. C. C. CRAB HOST : Cardisoma armatum. Indiscriminate breeder. Zatisquama (Coquillett. carcinoxenus Castro. Indiscriminate breeder. 1907 DISTRIBUTION: Tropical Africa. coastal (Atlantic only?) CRAB HOST: Not recorded. Normally breeds in bromeliads. ssp. bisulcatus (Coquillett. CRAB HOST : Not recorded. REFERENCES: Castro. CRAB HOST : Not recorded. Philippines. TYPE: Transient. Note: Probably adults only in crabholes or burrows. 19 15: 305. 195 1: 119. 1932 DISTRIBUTION: Brazil. Type series from bamboo joints. 1889 DISTRIBUTION: Southern and western Australasian Region. 1903 DISTRIBUTION : American Mediterranean Region. REFERENCES: Bruce-Chwatt and Fitz-John. consimilis Newstead. C. CRAB HOST : Ucides cordatus. 1928 :347. TYPE: Accidental. interior and coastal. TYPE: Transient. CRAB HOSTS: Cardisoma guanhumi. . corniger Theobald. REFERENCE: Dyar. coastal and interior. REFERENCE: Howard. interior and coastal. 1920: 253. Dalziel. Indonesia. C. 1962:pl. REFERENCE: Belkin. 1906) DISTRIBUTION : Tropical America. 1915:246.

1928 : 362. coastal. coastal. : Howard. 253. Indiscriminate breeder. 1952:27. C. REFERENCES Bonne and Bonne-Webster. TYPE: Specific or semispecific.1972 BURROWINGLAND CRABSOF THE WORLD C. TYPE: Transient. also breeds in ground pools and pot holes. Bionomics poorly known. Hogue and Wirth. REFERENCE: Wanson. inflictus Theobald. 1950: 239. 1925 : 189. REFERENCES: Bruce-Chwatt and Fitz-John. 578. Note: May be confused with decens. Indiscriminate breeder. 1901 DISTRIBUTION: Tropical Africa. Dyar. invidiosus Theobald. Pratt and Seabrook. 1928 : 249. 19 15: 262 (as eremita) . Note : A complex of species. CRAB HOST: Cardisoma armatum. 1928: 391. foliaceus Lane. Most records from crabholes. Dalziel. Knab. REFERENCES: Dyar. C. decens Theobald. 195 1: 119. C. Usually breeds in a wide variety of ground habitats as well as in containers. interior and coastal. REFERENCE: Wanson. 1935:576. 190 1 33 DISTRIBUTION: Ethiopian Region. Dyar and Knab. CRAB HOST: Cardisoma armatum. 1901 DISTRIBUTION : Tropical America. Note: May be confused with invidiosus. C. 578-579. interior and coastal. REFERENCE: Stone. CRAB HOST: Not recorded. 1901 DISTRIBUTION : Ethiopian Region. 1920: 25 1. CRAB HOST: Not recorded. interior and coastal. Howard. habilitator Dyar and Knab. TYPE: Transient. 1935:576. TYPE: Probably transient. Dyar and Knab. TYPE: Semispecific. CRAB HOST: Not recorded. ca tor) . 1968: 6. 1906 DISTRIBUTION: Antilles and Trinidad. TYPE: Transient. 1945 DISTRIBUTION : Brazil. CRAB HOST : Cardisoma armatum. interior and coastal. C. 1915: 327 (as extricator) . 1910: 868-869 (as extri- . Dunn. All records from crabholes. duttoni Theobald.

1935:576. Hill. 578-579. CRAB HOST: Cardisoma armatum... 1970: 49. C. TYPE: Transient. Usually breeds in foul ground pools and ditches and large artificial containers. coastal (Atlantic). 1929: 327. TYPE: Transient. CRAB HOST : Not recorded. pruina Theobald. Pratt et al. perfuscus Edwards. pipiens quinquefasciatus Say. CRAB HOST: Not recorded. 1914 DISTRIBUTION: Tropical Africa. 220 Theobald. General ground pool breeder. janitor No. CRAB HOST: Not recorded. 1960 :99. C. 578-579. TYPE: Transient. TYPE: Specific. sometimes found in crabholes. 1935 :576. C. interior? and coastal. interior and coastal. REFERENCES: Belkin et al. interior and coastal. 1959 DISTRIBUTION: Bahamas. nigripalpus 1905:406-407. Hanney. REFERENCE: Dunn. Wanson. 1901 DISTRIBUTION: Western and central Africa.34 CONTRIBUTIONS IN SCIENCE C. Usually breeds in ground pools. small islands. TYPE: Transient. philipi Edwards. REFERENCE: Wanson. 1823 DISTRIBUTION : Cosmopolitan. C. Branch and Seabrook. 1959: 216. Also found in vegetated pools. CRAB HOST: Not recorded. Vector of filariasis over wide areas of world. CRAB HOSTS: Cardisoma armatum. 1901 DISTRIBUTION : Tropical America. Martini. scimitar Branch and Seabrook. REFERENCE: Edwards. Type series bred from larvae found in crabholes. Not recorded from other sites. Sudanonautes africanus. 1903 DISTRIBUTION: Greater Antilles. C. C.. 1914:70 (as prasinopleurus). 1970:49. Grabham. 1928 :249. REFERENCE: Belkin et al. interior and coastal. TYPE: Transient. 1929 DISTRIBUTION: Western tropical Africa. CRAB HOST: Not recorded. coastal. Usually breeds in ground pools with decaying leaves. 1928 :249. Hill and Theobald. 1948 : 55. . REFERENCES: Dunn. 1945:246.

Also breeds commonly in saline ground pools and artificial containers. coastal. Ingram and Macfie. TYPE: Transient. TYPE: Semispecific. spathif urea (Edwards. 1828 DISTRIBUTION: Oriental Region. CRAB HOST: Not recorded. 1929:327. 1946 DISTRIBUTION: New Guinea. Strongly anthropophilic. A general ground pool breeder. 1959: 2 16. REFERENCE: King and Hoogstraal. REFERENCE: Branch and Seabrook. C. 1946b. interior and coastal. 1920:251-253. C. 578-579. TYPE: Semispecific or transient. nailoni King and Hoogstraal. . 1935:578. REFERENCE: van Someren et al. 1922 DISTRIBUTION: Ethiopian Region. 1952:286. 1917: 147-149. CRAB HOST: Cardisoma armatum. Wanson. C. Immatures unknown. interior and coastal. Wanson. thalassius Theobald. CRAB HOST: Not recorded. REFERENCES: Bruce-Chwatt and Fitz-John. 1955:487. Subgenus Culiciomyia C. CRAB HOST: Not recorded. Sesarma africanum. Dalziel. 195 1: 119-l 20. captured at the entrance of small crabholes in partial shade on the beach. REFERENCE: Steffan. east Africa and western Pacific. “crabhole in rain forest. CRAB HOST: Not recorded. Uca tangeri. interior and coastal. ruthi Peters. Numerous records from crabholes. REFERENCES: Dalziel. 1966:219. 1948: 145 (as stylifurcatus). 1935 :576. 1928: 249. Adults only.. cinerellus Edwards. C. coastal. Normally breeds in saline or brackish coastal ground waters. 1958 DISTRIBUTION: New Guinea. Dunn. Edwards. 19 15) DISTRIBUTION: Oriental and Indomalayan Regions. Multiple records from crabholes. Anthropophilic. C. REFERENCE: Carter and Wijesundara. TYPE: Transient. CRAB HOST: Cardisoma armatum. Adults only.1972 BURROWINGLAND CRABSOF THE WORLD 35 TYPE: Transient? Bionomics poorly known. Hopkins. One record from crabhole. sitiens Wiedmann. recorded also from ground pools. TYPE: ? Bionomics insufficiently known. coastal.” TYPE: ? Bionomics insufficiently known. 1903 DISTRIBUTION: Tropical Ethiopian Region. 1920: 25 l-253 (as nebulosus) .

: C. CRAB HOST: Not recorded. primarily coastal (?) CRAB HOST: Not recorded. reidi. variatus (Leicester. Japan. SubgenusLutzia C. interior and coastal. small islands. pl. TYPE: Transient. C. TYPE: Transient. Known only from crabholes. 1957:29. infant&us Edwards. REFERENCE:Macdonald. Usually breeds in shadedpools at margin of tidal zone. 1962 DISTRIBUTION : Solomons. 1908) DISTRIBUTION Indomalayan Region. interior and coastal. rubithoracis (Leicester. . coastal. TYPE : Transient. A general ground pool and container breeder.Collected repeatedly and exclusively from small crabholes in secondary rain forests in mountainous terrain. CRABHOST: Not recorded. interior. tigripes Grandpre and Charmoy. pholeter Bram and Rattanarithikul.36 CONTRIBUTIONS IN SCIENCE No. : CRAB HOST: Not recorded. REFERENCE:Belkin. coastal. 1901 DISTRIBUTION Ethiopian Region. 1962: 266. : C. CRAB HOST: Not recorded. 1967 : 13. TYPE: Specific. 1965: 273. 1908) DISTRIBUTION: Indomalayan Region. : CRABHOST: Not recorded. TYPE: Transient. Colless. 220 SubgenusLophoceraomyia C. C. REFERENCE:Colless. REFERENCE Bram and Rattanarithikul. Usually breeds in ground pools but also utilizes containers near the ground. 1965:280. REFERENCE:Colless. 2. Ground pool breeder. REFERENCE Bram. becki Belkin. 1967 : 6 1. 1967 DISTRIBUTION: Thailand. 1965 DISTRIBUTION Singapore. 1922 DISTRIBUTION : Oriental (including Japan) and Indomalayan Regions. C. : CRAB HOST: Cardisoma armatum. Selangor. TYPE: Specific.

coastal. interior and coastal. Probably normal breeding site deep seepage channels or solution holes in coral. Montchadsky and Garcia. TYPE: Accidental. inconspicuosus Duret. in streams and Iin ground pools. . 1966: 133. interior and coastal. 1968:41 (as cedecei) . CRAB HOST: Cardisoma guanhumi. REFERENCE: Wanson. 1967 : 80. CRAB HOST: Cardisoma armatum. 1967 (Theobald. 1906: 220. 254. TYPE : Specific. Usually breeds in nearly stagnant pools. 1935 DISTRIBUTION: Philippines. 1969 (T). REFERENCES: Pratt et al. carcin’ philus o Dyar and Knab. 1906 DISTRIBUTION: Dominican Republic. REFERENCES: Dyar and Knab. 1918 DISTRIBUTION : Tropical America.. REFERENCE: Dalziel. opisthopus 37 Komp. 1966:46. laureli Baisas. Indiscriminate breeder. 1926 DISTRIBUTION: Tropical America. REFERENCE: Delfinado. REFERENCE: Pratt and Seabrook. TYPE: Transient. 1952:27. nicaroensis DISTRIBUTION : Cuba. Belkin. TYPE: Transient. Stone and Hair. ? Guatemala. CRAB HOST: Not recorded. 1935 :578. Larva predaceous. Subgenus Mochthogenes C. Usually breeds in coastal ground pools shaded by mangroves. 1920:251-252. Subgenus Melanoconion C. C. CRAB HOST : Not recorded.1972 BURROWINGLAND CRABSOF THE WORLD TYPE: Transient. Cuba. C. coastal (Atlantic). C. iolambdis Dyar. coastal. 1945:246. 1908) DISTRIBUTION: Ethiopian Region. TYPE: Transient. C. CRAB HOST: Cardisoma guanhumi. CRAB HOST: Not recorded. TYPE: ? REFERENCE: Duret. primarily coastal (Atlantic). Usually breeds in small vegetated ground pools.

Multiple records only from crabholes. interior and coastal. 1958:90. 1970). rima Theobald. 1971. Referred to as “Crabhole Mosquitoes” in the literature they unquestionably are specific members of the crabhole community. CRAB HOST: Sudanonautes africanus. TYPE: Specific or semispecific. 190 1 DISTRIBUTION : Ethiopian Region. Provost and Haeger. 1960: 99. Deinocerites normally breeds in crabholes. 1928:249. Dalziel. Belkin and Hogue. Hanney. in spite of the occurrence of suitable habitats and hosts (Cardisoma and Ucides) along the entire Brazilian coast to Sao Paulo. 1959. mangrove pot holes and coastal ground pools. Macfie and Ingram. . Genus Deinocerites GENERAL REFERENCES: Adames. C. 19 16: 11. TYPE: Transient. TYPE: Transient. Philip. being found breeding outside this habitat only very rarely in such related or proximate places as mangrove treeholes. CRAB HOST: Cardisoma armatum. C. 1901 DISTRIBUTION: West and central Africa. some are definitely functional in their relationship to the crabhole habitat and community. REFERENCES: Dalziel. each species having a completely Atlantic or Pacific (with a few exceptions) distribution. 1920:251-253. interior and coastal. 220 (Carter. amphibian or bird blood (Templis and Galindo. interior and coastal. CRAB HOST: Cardisoma armatum. Cardisoma armatum. Dunn. 1959 :421) . All stages exhibit unique characteristics among the Culicidae. REFERENCE: Dalziel. 578. Usually breeds in forest ground pools and streams. This apparent truncation in the distribution may be due only to lack of collecting. the most southernly Atlantic record is on the coast of the State of Maranhao (Cerqueira. 1920: 251-253. 1920:251-253. Wanson. REFERENCES: Bruce-Chwatt and Fitz-John. 1911) DISTRIBUTION : Africa. Usually breeds in foul water in pooled streams. Only occasional specimens are observed biting man. The genus ranges throughout the American tropics. 1935:576. Curiously. salisburiensis Theobald. 1931: 192. The peculiar pupal attendance and mating behavior first described in detail in Deinocerites cancer (Downes. 1938: 291). 1966.38 CONTRIBUTIONSIN SCIENCE Subgenus Neoculex C. insignis No. Surtees. 195 1: 119. The normal food is probably reptile. Among the immatures the larvae of all species have lateral head pouches of unknown function and the pupae of some have three float hairs (these and others discussed by Belkin and Hogue. 1967) represents an adaptation correlated with a non-dispersing evolutionary trend in this line of mosquitoes.

Peyton et al. 1966 (B) . D. barretoi Adames. Komp. 1907 CRAB HOST: Not recorded. D. D. melunophyllum Dyar and Knab. subcylindrica. 1956. mathesoni Belkin and Hogue. 1971 CRAB HOST: Not recorded. D. REFERENCES: Fisk. D. costaricensis Adames and Hogue. 1901 CRAB HOST: Cardisoma guanhumi. 197 1 CRAB HOST: Not recorded. CRAB HOST : Not recorded. Known from small crabholes. 1970 CRAB HOST : Ucides occiden talis. REFERENCES: Downes. 1959)) is known to be autogenous. colombianus Adames.1972 BURROWINGLAND CRABSOF THE WORLD 39 Deinocerites cancer. U. mcdonaldi Belkin and Hogue. cancer Theobald. nicoyae Adames and Hogue. 1970 CRAB HOST: Cardisoma crassum. 1959 CRAB HOST: Not recorded. D. 197 1 CRAB HOST: Not recorded. at least in Florida (Haeger and Phinizee. belkini Adames. D. atlanticus Adames. Provost and Haeger. Uca (?) D.. D. epitedeus (Knab. D. 1959 CRAB HOST: Not recorded. . magnus (Theobald. 1901) 1959 (B) . 1959 CRAB HOST: Not recorded. 1907) CRAB HOST: Not recorded. 197 1 CRAB HOST: Not recorded. curiche Adames. D. howardi Belkin and Hogue. D. D. 1959 CRAB HOSTS: Uca pugilator. D. dyari Belkin and Hogue. D. 1967 (B). Haeger and Phinizee. 1941 (as spanius) (B) . 197 1 CRAB HOST : Not recorded. Gecarcinus lateralis. 1964 (B).

Apparently normally breeds in empty Achatina shells (land snails). Peyton (Dyar and Knab. 1935 :576-577. Adults only. 579. 220 Adames. TYPE: Probably specific. probably breeds in vegetated jungle pools like its relatives. aurites (Theobald. TYPE: Accidental. guanhumi. 1909 CRAB HOSTS: Cardisoma Gecarcinus lateralis. interior and coastal. Genera Mansonia and Coquillettidia All occurrences of Mansonia and Coquillettidia are resting adults only and constitute accidental utilization of the crabhole habitat. Genus Eretmapodites E. quinquevittatus Theobald. C. CRAB HOST: Not recorded. CRAB HOST: Cardisoma armatum. CRAB HOST: Cardisoma armatum. D. Genus Galindomyia G. spanius 1967 (B) . TYPE: Accidental. 190 1) DISTRIBUTION: Tropical Africa. The larvae of all members of these genera are associated strictly with floating and emergent water plants from which they obtain oxygen with a specially modified. pseudes Dyar and Knab. 1935 :576. REFERENCES: Galindo. . D. though adults only known from crabholes. CRAB HOST: Cardisoma armatum. Note: Anthropophilic. 1968. REFERENCE: Wanson. C. REFERENCE: Wanson. Hogue and Wirth. crassurn. panamensis No. 190 1 DISTRIBUTION: Tropical Africa. Uca subcylindrica. piercing siphon. 1909) CRAB HOST: Not recorded. 1969 DISTRIBUTION: Colombia. 197 1 CRAB HOST: Not recorded. REFERENCE: Wanson.. 1969. Genus Hodgesia H. interior and coastal. Biology not known. interior and coastal. leei Stone and Barreto. REFERENCE: Stone and Barreto. 1930 DISTRIBUTION: West tropical Africa. 1964 (B).40 CONTRIBUTIONSIN SCIENCE D. 1935:576. et al. coastal. nigeriae Edwards.

confinnis (Lynch Arribalzaga. Sudanonautes africanus. interior and coastal. 1961:25. 1952:59. 1960 : 99. interior and coastal. U. Note: Anthropophilic and vector of yellow fever and filariasis. Many records and collections. CRAB HOSTS: Cardisoma arma turn. Usually breeds in plant and artificial containers and ground pools and in sago palm swamps. Hopkins. alboabdominalis Theobald. CRAB HOST: Cambarus diogenes ludovicianus. TYPE: Transient. 1962. CRAB HOST: Not recorded. Macfie and Ingram. . CRAB HOST: Cardisoma armatum. 1920 :253. See Peyton. CRAB HOST: Cardisoma armatum. 1931: 192. 1958:91. REFERENCE: Dalziel. Caribbean. 1928 :249. coastal. Dunn. TYPE: Transient. Steffan. REFERENCE: Wanson. REFERENCES: Bruce-Chwatt and Fitz-John. REFERENCES: van den Assem. interior and coastal. 1966:203. 1905 DISTRIBUTION: New Guinea. atra Theobald. 19 10 DISTRIBUTION: Tropical Africa. Bismark Archipelago. Surtees.1972 BURROWINGLAND CRABSOF THE WORLD M. Hanney. Dalziel. eastern South America to Argentina. Usually breeds in shallow ground pools. TYPE: Transient. 195 1: 119. 253 (as fasciata) . U. 1901) DISTRIBUTION: Tropical Africa. 1960 : 99 . A ground pool breeder. Philip. africana (Theobald. U. uniformis (Theobald. TYPE: Specific or semispecific. 1920: 251. all from crabholes. 1916:7. Note: Anthropophilic and important vector of filariasis. CRAB HOST: Sudanonautes africanus. annulata Theobald. 189 1) DISTRIBUTION: Eastern and southern United States. Genus Psorophora P. Genus Uranotaenia Nearly all the records in this genus are from the burrows of freshwater crabs. REFERENCE: Hanney. interior and coastal. interior and coastal. Adults only. Note : A complex of species. REFERENCE: Evan. 1970. 190 1) 41 DISTRIBUTION: Widespread throughout Old World Tropics. M. 1901 DISTRIBUTION: Western tropical Africa. 1935 :576.

TYPE: Accidental. Thailand. 1908 :8. 220 U. 1908 DISTRIBUTION Indomalayan. REFERENCE Leicester. REFERENCE:Peyton and Klein. REFERENCE:Qutubuddin. 1908 DISTRIBUTION Southeast Asia. : CRAB HOST: Not recorded. U. CRABHOST: Not recorded. CRAB HOST: Sudanonautes africanus. interior and coastal. 190 1 DISTRIBUTION: Tropical Africa. koli Peyton. : CRABHOST: Not recorded. 19 12 : DISTRIBUTION Tropical Africa. Adults only. 193 1 DISTRIBUTION: Nigeria. Bionomics insufficiently known. : U. TYPE: ? REFERENCE:Philip. etc. TYPE: ? Adults only. mashonaensis Theobald. Usually breeds in slightly brackish open pools behind beaches. interior. 1970: 3 and personal communication. REFERENCE:Hanney. U. although may be very common in crabholes. U.). 1970 DISTRIBUTION Cambodia. 1960: 99. TYPE: Accidental? Probably normally breeds in grass-grown ground pools. TYPE: Transient. 2 17 (as cancer). Fresh water species. fraseri Edwards. Usually breeds in a wide variety of ground pools. 1946 ( 1947) DISTRIBUTION India. 1920:251-253. U. Madagascar. bicolor Leicester. pools. interior and coastal. interior.42 CONTRIBUTIONS IN SCIENCE No. rockpools. interior. 1946 ( 1947) : 118. TYPE: Specific or semispecific. Breeds in various ground habitats (swamps. husaini Qutubuddin. CRAB HOST: Not recorded. : coastal. REFERENCE:Dalziel. U. TYPE: Transient. caliginosa Philip. 1970: 248. north Australian and Papuan Regions. bilineata var. REFERENCE:Peyton. . : CRABHOST: Not recorded. lateralis Leicester. interior. 193 1: 190. CRABHOST: Not recorded.

nivipous Theobald. CRAB HOST: Not recorded. 1943. 195 1: 107. Hyderabad (Deccan) City. .” REFERENCE: Qutubuddin. coastal? or interior by large rivers? CRAB HOST: Not recorded. Cape Province. montana Ingram and de Meillon. TYPE: ? Adults only. 1966 DISTRIBUTION: Philippines. Transvaal. coastal. TYPE: ? Bionomics insufficiently known. usually being encountered in fresh water ponds and lakes. 1970: 3 and personal communication. 1958:91 (as candidipes). U. Adults only. 195 1 43 DISTRIBUTION: India. India. TYPE: ? Bionomics insufficiently known. 1912 DISTRIBUTION: Tropical Africa. REFERENCE: Hopkins. taken at mouths of crabholes. philippinensis Delfinado. CRAB HOST: Cardisoma sp. CRAB HOST: Not recorded. U. Genus Corethrella C. Numerous collections all from crabholes. Family CHAOBORIDAE The larvae of phantom midges are all aquatic. Bionomics insufficiently known. in October. stonei Lane. “Habitat: caught from crabholes in an old pond in the Public Garden. (guanhumi?) TYPE: ? Bionomics insufficiently known. Those of two species in the genus Corethrella are recorded from crabholes. 1942 DISTRIBUTION: Panama. A single record from crabholes. 1927 (as cundidipes) . REFERENCE: Lane. REFERENCES: Ingram and de Meillon. rossi Delfinado. interior. interior. interior. U. 1952 :58. coastal? CRAB HOST: Not recorded. Two records only from crabholes. TYPE: Specific or semispecific. REFERENCE: Peyton. REFERENCE: Peyton.1972 BURROWINGLAND CRABSOF THE WORLD U. U. 1970: 3 and personal communication. Surtees. 1966 DISTRIBUTION: Philippines. The larva recorded by Hopkins was collected from a crabhole and lived in captivity for 4l/2 months before pupating. 1942: 119. TYPE: ? Bionomics insufficiently known. 1927 DISTRIBUTION : Natal. CRAB HOST: Not recorded. mattinglyi Qutubuddin.

19 13 DISTRIBUTION: Mesoamerica. Gold Coast. interior coastal. coastal? CRAB HOST: Not recorded. TYPE: ? Bionomics insufficiently known. 1942 : 120. CRAB HOSTS: Cardisoma crassurn. CRAB HOST: Not recorded. D). Genus Culicoides C. 1956:197 (B. Wirth and Arnaud. 1944: 109. Usually breeds in coastal marshes. Family CERATOPOGONIDAE A few species of this family. 100 feet elevation. CRAB HOST: Cardisoma armatum. 584 (as wansoni). arubae Fox and Hoffman. REFERENCE: Fox and Hoffman. salt marsh and tidal flat-loving flies will be found utilizing the crabhole habitat. C. TYPE: Transient. Uganda. TYPE: Specific. 1969 :5 17-5 18. and 1944 .D). Forattini et al. CRAB HOST: Not recorded. 220 DISTRIBUTION: Trinidad. REFERENCE: Hogue and Wirth. are recorded as breeding in crabholes. cancer Hogue and Wirth. Ucides occidentalis.44 CONTRIBUTIONSIN SCIENCE C. coastal. TYPE: ? Bionomics insufficiently known. 1946 DISTRIBUTION: Fiji Islands. REFERENCE: Wanson. 1968 DISTRIBUTION : Costa Rica.. all in the genus Culicoides. REFERENCES: Macfie.. DISTRIBUTION: American Tropics. 1946. Senegal. tripunctata Lane. coastal. 19 12 DISTRIBUTION : Nigeria. CRAB HOST: Ucides cordatus. REFERENCE: Lane. C. Larvae found in crabholes beneath rocks. distinctipennis Austen. Brazil. C. TYPE: ? Bionomics insufficiently known. 1968 (G). 1942 No. Certainly many more of these coastal. insignis Lutz. The species has been collected in all stages numerous times and in very large numbers only from crabholes. C. 1958: 37 (B. cancrisocius Macfie. coastal (Pacific). TYPE: ? Bionomics insufficiently known. 1935:579. coastal. Puerto Rico. REFERENCES: Forattini et al.

Others (species largely unidentified) are known in the immature stages from water in the burrow. Numerous collections. REFERENCES: Carson. The larvae live on the crabs and even pupate (in one species) on the third maxilliped. usually as adults only. Family CHIRONOMIDAE Species undetermined.D). Nigeria. Congo. small islands. D. 19 13 45 DISTRIBUTION: Brazil. 1967 (B. it probably consists of food leavings of the host in one case and the host’ tissues in the other.D). MISCELLANEOUS DIPTERA A number of shore inhabiting flies have been taken. 1960 (B.D). all from crabholes.D) . Though the food of the larvae is not known with certainty. coastal. TYPE: Parasitic. Banana. carcinophila Wheeler. Lutz. from the mouths of crabholes. CRAB HOST: Cardisoma guanhumi. Bahamas. running over and hovering about the carapace. Panama. TYPE: Commensal. islands only (coastal on large islands). 1960 DISTRIBUTION: Greater and Lesser Antilles. 1968 (B. Wanson. TYPE: Specific or semispecific. REFERENCES: Forattini et al. s Genus Drosophila D. Wheeler.1972 BURROWINGLAND CRABSOF THE WORLD C.. 195 1: 118. The adult flies have been observed to remain on the crab. LOCALITIES: Lagos. reticulatus Lutz. endobranchia Carson and Wheeler. CRAB HOST : Gecarcinus ruricola. Providencia. 1912:19. 1968 DISTRIBUTION : Cayman Islands. TYPE: ? REFERENCES: Bruce-Chwatt and Fitz-John. Family DROSOPHILIDAE Two species of Drosophila have developed a symbiotic association with land crabs of the genus Gecarcinus. larva on host’ gills. s REFERENCE: Carson and Wheeler. CRAB HOST: Cardisoma armatum. 1957:312 (B. 1935: 578. . 1913: 50 (B). larva in renal grooves and peribuccal region. CRAB HOSTS: Gecarcirzus rzlricola and lateralis.

Undetermined “larva A” LOCALITY: Admiralty Islands.46 CONTRIBUTIONS IN SCIENCE No. TYPE: ? REFERENCE: Wheeler. 1938: 126-129. REFERENCES: Baylis. 1960:210. Adults rest and hide in the burrow mouth. REFERENCES Baylis. Immatures undoubtedly develop elsewhere. coastal. Not recorded. Keilin. REFERENCE: Melander. LOCALITY: Not specified. REFERENCES: Wheeler. CRAB HOST: Not recorded. CRAB HOST: Ocypode ceratophthalma. 1938 DISTRIBUTION: Hawaii. TYPE: Symbiosis ? Larvae found in branchial chambers of preserved crabs. Not specified. Keilin. Family DOLICHOPODIDAE Asydetus carcinophilus Parent. 19 15. 1938:57. : chambers of preserved . 19 15. TYPE: ? REFERENCE: Wheeler. CRAB HOST: Cardisoma hirtipes. 1960: 210. 1960:210. Undetermined “larva B” LOCALITY: Christmas Island (Indian Ocean). Family EPHYDRIDAE Hecamede sp. 1937 DISTRIBUTION: Hawaii. TYPE: Transient. CRAB HOST: Not recorded. CRAB HOST: Gecarcoidea lalandii (humei). Williams. probably Ocypode. TYPE: Symbiosis ? Larvae found in branchial crabs. Adults found on the beach in the near vicinity of burrows. coastal. 220 Family CHLOROPIDAE Genus Lasiopleura LOCALITY: CRAB HOST: sp. 192 1. 192 1. Family EMPIDIDAE Chersodromia hawaiiensis Melander. TYPE: Transient.

Primarily a flatwoods species occurring in slow streams. TYPE: ? REFERENCE: Wheeler. 1941 47 DISTRIBUTION: Florida. ponds. A streamside mud flat inhabitant. REFERENCE: Hanney. 1952 DISTRIBUTION : Costa Rica. interior. 1960 : 100 (as Legnotus tibialis) . Family CYDNIDAE Sehirus tibialis Stal. MISCELLANEOUS INSECTS Order HEMIPTERA Family GELASTOCORIDAE Mononyx grandicollis Germar. interior and coastal? CRAB HOST: Sudanonautes africanus. ditches. 1840 DISTRIBUTION: West Africa. interior and coastal? CRAB HOST: Sudanonautes africanus. TYPE: Transient or accidental. 1960: 100. . TYPE: ? REFERENCE: Drake. Order COLEOPTERA Family DYTISCIDAE Bidessus rogersi Young. TYPE: Transient. TYPE: Accidental. CRAB HOST: Not recorded. and other lenitic situations. 1952: 14-15. CRAB HOST: Not recorded. REFERENCE: Hogne and Wirth. cypress swamps. 1968: 6. Larva only. CRAB HOST: Procambarus rogersi rogersi. REFERENCE: Young. Ground burrowing species. 64-65. REFERENCE: Hanney. TYPE: ? Bionomics insufficiently known. 1960: 2 10. CRAB HOST: Cardisoma crassurn. 1853 DISTRIBUTION: West Africa.1972 BURROWINGLAND CRABSOF THE WORLD Family TETHINIDAE R hicnoessa sp. 1954: 17. Family VELIIDAE Microvelia oraria Drake. Family HELODIDAE Helodes ? sp. LOCALITY: Costa Rica. LOCALITY: Not specified.

Key West. CRAB HOST: Gecarcinus lateralis. Family LAELAPTIDAE Laelaps cancer Pearse. 1929 :229-230. DISTRIBUTION: Dry Tortugas Islands. 1967 : 342. s REFERENCES: Pearse.48 CONTRIBUTIONSIN SCIENCE Order ACARINA No. Banana. certain other aquatic crustacea appear to be members of the crabhole community. Congo. small islands. REFERENCE: Pearse. TYPE: ? REFERENCES: Bruce-Chwatt and Fitz-John. 1935 : . Nigeria. Class CRUSTACEA Apart from the crab host itself. 19 13 :264-268. coastal. All stages found in branchial chambers and on gills. TYPE: Commensal or parasitic. Order EUCOPEPODA Cyclops sp. Probably feed on host’ blood or secretions. CRAB HOST: Gecarcinus ruricola. TYPE: ? REFERENCE: Carson. Wanson. Single nymphal specimen known. 1929 :230. 578. 195 1: 118. TYPE: Parasitic? Cling to gill filaments of crab with their second antennae and maxillipeds. REFERENCES: Pearse. small islands. 1929 DISTRIBUTION: Dry Tortugas Islands. CRAB HOST: Gecarcinus lateralis. 1932 : 112. Wilson. 220 Two mites of different families are known as symbiotic associates of certain land crabs : Family TYROGLYPHIDAE Rhizoglyphus sp. Family UNDETERMINED Species Undetermined LOCALITY: West Indies. 1932: 112. CRAB HOST : Cardisoma guan humi. TYPE: Commensal or parasitic. Order COPEPODA Cancrincola janzaicensis Wilson. CRAB HOST: Cardisoma armatum. LOCALITIES: Lagos. 19 13 DISTRIBUTION : Jamaica.

Training Manual. and Dr. including Ceylon and Burma. R. University of California. Bull. J. K. London XIII. 1962. A parasitic oligochaete and other inhabitants of the gillchambers of land crabs. John N. Press. University of Southern California. 337 p. J. Ser. Natur. L. Berkeley. Insects injurious to man and stock in Zanzibar. Mag. Los Angeles County Museum of Natural History Foundation. S. Res. Tribes Megarhinini and Culicini. Louisiana. E. 6. Descriptions of new species of South African Decapod Crustacea. David Grajeda. Natur. BEHRE. Descriptive catalogue of South African Decapod Crustacea. LITERATURE CITED ADAMES. Mus. Materials for a carcinological fauna of India. 15:378-381. Fullerton. BELKIN. Amer. Peyton and Botha de Meillon. 1970. 1947. Tiere 48: 123-146. ALCOCK. S. Univ. Syst. Mosq. J. PAGE. 20). Mosq. University of Queensland. 197 1. Zeit. 22 pl.. 1900. BAYLIS.1972 BURROWINGLAND CRABSOF THE WORLD ACKNOWLEDGMENTS 49 For direct assistance and contributions to the preparation of this paper the authors wish especially to thank the following individuals: Drs. with notes on synonomy and new records. _. E. William Macnae. Hist. We are also indebted to many institutions and persons who have offered assistance and guidance in connection with our field trips through Central and South America. The mosquitoes of the South Pacific. Inst. 1917. Sond. Most significantly these are: the American Philosophical Society. C. Contrib. Jocelyn Crane. Okol. J. Los Angeles. 1915. N.. AMOS. A. 6( 1): l-458 (also Inst. H. Garth. ____ _. Hist. Ser. Asiatic Sot. Entomol. A revision of the crabhole mosquitoes of the genus Deinocerites. Okologische und ethologische studien an Europas einziger Winkerkrabbe Uca tangeri Eydoux. Dr. Mag. Griffin. 7: 1-154.. University of Witwatersrand. Entomol. Hist. 1934.. John S. Southeast Asia Mosquito Project. Shivaji Ramalingam. and William Stephenson. BARRAUD. from which were derived the original portions of this paper. Bengal 69:279-456. 1949. The Brachyura Catometopa or Grapsoidea. The fauna of British India. New York Zoological Society. 1944. W. Land crustacea. 11. ALTEVOGT. W. 38:1-837. H. . D. D. J. Deutschen Zool. 1950. Suva. ADERS. Culex (Melanocanion) annulipes invalid. Richard Dwyer. 7:391-401. Morph. Newsletter 1:68. Amer. A. Afr.. Belkin. 608 p. Ges. Louisiana Acad. Akustische epiphanomene im sozialverhalten von Ucu tangeri in Sudspanien. Inst. Jamaica. Bull. 1959. of Calif. Family Culicidae. No. M. Ann. The Culicidae of Jamaica p-_) (Insecta. D. BARNARD. National Geographic Society. 13:361-392. Martin. J. W. British Museum Natur. Verh. P. Diptera 5:1-463. and Mr. 1900. 12: 19-22. ANDREWS. Sci. Fiji. 43 p. 8. Ann. and W. Ann. HEINEMANN. Entomol. Ser. Allan Hancock Foundation. H. Australian Museum. Proc. + 412 pl. Wien 22:309-315. East Africa and Australia. Mosquito studies (Diptera. Notes on the occurrence of Cardisoma guunhumi Latreille at Grand Isle. A. Purvis L. Culicidae) XXIV. Control. California State College. . in A monograph of Christmas Island (Indian Ocean).-_. Diptera). A. Sci. University of Malaya. 1962. Contrib. 1969.

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