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When discussing anatomy of any animal, it's important to first gain a point of reference and establish terminology for

description. We can't use terms like "over" or "under" or "in front" or "behind." It's fine to say that the trachea is in front of the esophagus when we are looking at a penguin from the front, but the esophagus is in front of the trachea if we look at the penguin from behind. Vessels can run under a tissue from one point of view or over a tissue from another point of view. Therefore, to be consistent, we establish terms that apply to location from any point of view. Look at the following drawings to learn the anatomical planes.

The Adelie penguin on the left of the screen is bisected by three planes. You can see a plane running parallel to the ground, running right through the abdomen of the penguin. This is called the horizontal or transverse plane. There isn't just one horizontal plane; there can be many horizontal planes. However, all horizontal planes are parallel to the ground. The horizontal plane divides the penguin into top and bottom portions. I'll go into more specific terms for "top" and "bottom" a little later. This is called the axial plane as well

The next plane which is visible appears as a rectangle running around the penguin. This plane runs perpendicular to the ground and divides the penguin into front and back portions. This is called the frontal or coronal plane. Like the horizontal plane, there can be many coronal planes dividing up the penguin.

The last plane appears as a thin line running through the center of the penguin. This is called the median plane or sagittal. There is only one median plane and only one sagittal plane since by definition the "median" is the absolute middle. The sagittal plane is named after the mythological creature Sagittarius whose arrow would pierce victims and split them directly into two equal halves. Planes bisecting the penguin to the left or right of the median plane are called parasagittal planes. Why are these planes useful? Well, they are useful when we talk about parts of the penguin. For example, a vessel may run in the median of the penguin. When I say that, you might imagine that the vessel runs directly down the middle of the penguin. What if I say the nerve runs horizontally under the rib? You might imagine that this nerve runs in the horizontal plane. So, the purpose of planes is to give a frame of reference. Planes are only so useful. The terms that are more useful are the directional terms that indicate the position of organs or tissues. Look that the image below for some basic anatomical terms.

First let's talk about the terms dorsal and ventral. Dorsal is what we think of when we think of the back of the penguin. However, we can't say that the erector muscles of the back are "back to" the spinal column. But, we can say that the erector muscles are dorsal to the spinal column. (We can also say "superficial" or "posterior" to in this case.) So, dorsal refers to the back. If dorsal refers to the back, then ventral must refer to the front or "belly" of the penguin. So, if I say the nerve runs along the ventral aspect of the penguin, you might imagine that the nerve runs along the "bellyside" of the penguin. The second terms we use are cranial and caudal. Cranial means "towards the brain or cranium." Caudal means "towards the tail." Cranial is sometimes used interchangeably with superior, and caudal is sometimes used interchangeably with inferior. So, if I say that the foot is caudal to the knee, you know that the foot is anatomically "below" the knee. This is a ridiculous example, but you get the idea.

The last three terms are a little more confusing. First, the term rostral means "towards the nose." This doesn't seem too confusing, but sometimes rostral is used to mean "towards the head," which you know is "cranial." To understand which they mean one has to read closely. The last terms to know are proximal and distal. They are used to clarify "origin" and "termination." So, a structure that is proximal to another structure is a structure that is closer to the origin. Look at the flipper in the drawing. The "axilla" is the "armpit" of the penguin. Would you say that the axilla is proximal or distal to the tip of the flipper? Although it is far away from the tip of the flipper, anatomically, it is proximal to the tip of the flipper. "Proximal" and "distal" can be confusing terms, but all you need is a little practice. So let's dive right into penguin anatomy, and we'll start with penguin embryology. Penguins start somewhere, you know...

When we encounter penguins, we see mostly the external features of the penguin: the feathers, eyes, wings, feet, etc. We'll begin the detailed anatomy section by identifying the numerous and basic superficial features. In this case, all the superficial features are external features. External features are very important for penguins. Although the primary means by which penguins differentiate each other is by vocalization, there are many external features which provide clues to penguins. These clues contain information regarding age, speciation, sex and many more. Look at this Yellow-eyed penguin. There are many characteristics of the external body which give us clues when we want to differentiate; they probably provide similar means for other penguins.

The above image shows some of the features that we see when we look at a penguin. Most often we recognize the facial features of the penguin above all else, so when we talk about feathers that grow from the head of the penguin, now we can speak about them accurately. Some of these terms should be immediately obvious to us like crest, crown, forehead and chin. The two terms I want to elucidate are the auriculars and the jugulum. The auriculars are the feathers that cover the auditory canal on penguins. The jugulum is the most ventral portion of the throat. Remember these are all external features. The eye is one of the most dynamic features of the penguin, and there are several features that come in intimate contact with the eye externally. Primarily these refer to the features and skeletal ridges around the eye. The commissure is composed of skin that bridges the upper and lower mandible. All penguins have this structure, and it is varied in color across the species. Those are the primary facial features you should be able to identify in penguins. The rest of the features are bodily and they can be seen in the below image. They are useful when we talk about injuries to penguins or when we talk about portions of the penguin that are used in behaviours like the bill-to-axilla. We can also use these basic terms to describe plumage as well.

Regions of the penguin are defined by the terms breast, abdomen (or belly), flank (the lateral portion of the penguin caudal to the abdomen that meets the wing/flipper), and the rump. Feather portions noted in this image are the scapulars which cover the proximal flipper and the axilla and the crural feathers. "Crural" refers to the leg.

The biology system of the penguin is very complex, no less complex than our own. In fact, there are systems present in penguins that do not exist in mammals. There are several ways to analyze the biology of penguins, and the first place to start is at the cellular level. So that's where we'll begin. Bone The cardinal lesson that bone teaches is that the foundation of all endoskeleton creatures is that bone is living tissue. Bone consists of the following structures: Hydroxyapatite crystals Collagen fibers (type I collagen fibers) Osteocytes (bone cells) Osteoblasts (bone forming cells) Osteoclasts (bone remodeling cells) Blood vessels and nerves.

What are all these features of penguin bone? Hydroxyapatite is the amalgamation of calcium in bone. You've heard the expression that calcium builds bone. Well, calcium is the primary component of hydroxyapatite crystals, 40 um length depositions throughout calcified bone. Hydroxyapatite crystals constitute the bulk of the inorganic matrix of bone. The organic matrix of bone is composed primarily of collagen (Type I) fibers. Both these matrices are elaborated by osteoblasts and osteocytes. Osteoblasts begin the elaboration, and when osteoblasts become surrounded by what they've elaborated, the become known as osteocytes. Osteocytes secrete much less collagen than do osteoblasts. Osteocytes are the permanent living cells of the bone matrix. The final cell in bone is the osteoclast. Osteoclasts are what remodel bone. They are stimulated by Vitamin D on the surfaces of osteocytes. When so stimulated, osteoclasts break down bone and allow growth and remodeling to take place in penguins. The image to the left is a typical cross-section of bone stained with the standard hematoxylin and eosin stain. H&E provides the characteristic pink and red stains you might find in histological sections such as this. The dark pink web-link branching throughout the image you see on the left is decalcified bone. That is, the calcium/hydroxyapatite has been removed from this section through staining techniques. Punctuated along the decalcified bone are the osteoblasts. You can also see blood vessels (BV) running through the marrow cavity (MC). The presence of blood vessels in bone implies that bone is indeed a living tissue and that the cells require nutrients, especially oxygen. The marrow cavity is the site of blood cell proliferation. In the adult penguin, blood cells in the body stem from the marrow cavities. When bone elaborates in a genetically determined pattern, they form larger structures in penguins you are probably familiar with.

So, where do penguins get calcium to build their bones? And where do they get the Vitamin D they need? Penguins eat a lot of seafood, and fish liver oil contains a significant source of Vitamin D. Those same fish contain soft bones, and bones, as you know, are rich in calcium. To build their own bones, the small fish and crustaceans that penguins consume provide these necessary nutrients. Typical foods penguins eat to build their calcium and Vitamin D stores: Crustaceans Shoaling fish Squid Krill (Plankton)

The skeletal system is the basis of support for penguins. It binds muscles together and it provides protection for the internal organs. It allows penguins to stand upright and prevents them, basically, from becoming a ball of mush. Penguin skeletons are evolutionarily designed for flight, despite the fact that penguins don't fly. However, penguin bones are slightly more dense than the average avian skeleton. And if you've been to the histology section, you know that the skeleton is a living and dynamic system. I'm not going to enter a detailed discussion of muscular attachments, but I will point out the basic components of the penguin skeleton and how they are important to the daily functioning of a penguin. Exposure will be cranial to caudal. There is one important definition that I will develop at this point. Articulation: the point, area, etc. where one bone meets and interacts with another bone. (verb: articulate, articulates). The image above is obviously the skull of a penguin -- a King penguin to be exact. There are several features you should know about the basic penguin skull, and they are all easily visible on this picture. What you know as the bill or "mandible" is really composed of two separate portions. One is connected directly to the premaxilla; this is known as the maxillary process. The inferior part of the mandible is known as the dentary. Contained within the maxillary process is the nasal aperture; this is the site through which scents and air enter the olfactory system of the penguin. As we move down the list of terms on the first image, we come across three terms for portions of the skull: the frontal portion, the parietal portion, and the occipital portion. They correspond to portions or lobes of the penguin brain. The zygomatic arch is a thin strip of bone that forms the inferior base of the "eye socket." The mesethmoid is a sinus that is lined by the same tissue that lines the respiratory system -- pseudostratified columnar epithelium. The figure below demonstrates the fundamental aspects of the bony skeleton. Although the bill or beak appears to be part of the skeletal system, it is not. As you know, the penguin bill is a keratinous structure, not hydroxyapatite. Its thick and compact molecular structure allows it to persist. The bill is connected to the pre-maxilla of the penguin. The pre-maxilla forms a portion of the cranium. The cranium contains one of the more important organs of the penguin: the brain, the coordinating center of the penguin. Caudal to the skull are the cervical vertebrae. Cervical vertebrae do not directly articulate with the base of the skull, however.

They are united via several ligaments, which are dense, tough connective tissue. There are several cervical vertebrae; other than the first, each vertebra articulates with a vertebra above and below it. The cervical vertebrae quickly change morphology and become known as the thoracic vertebrae. Thoracic refers to the thorax or chest. This is where the "rib cage" begins. The head of each rib articulates with a thoracic vertebra. The numerous ribs form a protective shield for the internal organs. The most elaborate thoracic bone is the most ventral: the sternum and keel. Note that this bone is unusually large. The pectoralis muscles attach to the keel of the sternum and extend to the humerus of the upper limb. Why is this sternum so large? Birds fly, and in order to fly, they need particularly large and powerful pectoralis muscles. In the case of penguins, they do not fly; however, they do swim and also require large pectoralis muscles. Sterna and keels of penguins are large as well. The pectoralis is the primary depressor muscle, and it runs ventromedially to the keel from the humerus. The primary elevator of the penguin is the supracoracoideus which runs dorsomedially from the humerus to the articulation of the clavicle and the coracoid bone. (These muscles are analogous to the pectoralis major and the latissiums dorsi in human beings.) In penguins, this entire "flight" structure is known as the pectoral girdle. The limbs of the penguin are similar in skeletal structure, blood supply and muscular compartmentalization. Each of the extremities (limbs) is composed of a singular bone (humerus=upper limb; femur=lower limb) proximally, two intermediate bones (radius and ulna=upper limb; tibiotarsus and fibular=lower limb), metacarpals (upper limb) or metatarsals (lower limb) and phalanges distally. In penguins, one cannot usually see the femur (thigh) since it is buried below the feathers. The last structure is the pubis. The pubis of a penguin is an interesting structure when it is compared to the pubis of a dinosaur. That of the penguin points toward the head; that of the dinosaur points to the tail. This evolutionary change seen in birds (and some advanced dinosaurs) is called the reverse pubis.

Both bone and muscle require blood in order to live. That is to say, penguin tissues need oxygen and they are delivered by blood vessels. Blood vessels come in intimate contact with almost all parts of the penguin body. Let's take a closer look. Blood In order to get oxygen to a penguin's tissues, something needs to carry that oxygen. In mammals, these cells that carry oxygen are called erythrocytes. You probably know them as red blood cells since they look red when they are carrying oxygen (although the color of cells is ultimately dependent upon what dyes people use to stain cells when they look at them under the microscope. Take a look at the following slides of red blood cells. One slide belongs to a mammal, like us, and the other belongs to a penguin. Can you guess which is which? (HINT: the red blood cells are stained greenish-brown in the top slide and yellow in the bottom one).

Have an answer yet? Well, the top slide is of mammal blood and the bottom one is penguin blood. What do you notice immediately about penguin erythrocytes and mammal erythrocytes? (HINT: It's not the color of the stain.) If you noticed that penguin red blood cells have nuclei then you are correct! That's right; mammal red blood cells lose their nuclei late in their development. Penguin and all avian red blood cells retain their nuclei. (The other cells which are stained blue are, oddly, white blood cells or platelets.) So now you know about penguin red blood cells and that they carry oxygen. How are these and other blood cells carried through the penguin body?

A: Capillaries B: Superficial plexus C: Subsuperficial plexus D: Intermediate plexus E: Deep plexus

Larger blood vessels give rise to smaller blood vessels, and eventually these give rise to capillaries. As you can see, capillaries are the end of the trip oxygen takes in the blood. When red blood cells reach the capillaries, they travel one-by-one, in single file through those capillaries to deliver their oxygen. Capillaries wind in and out of muscle fascicles and bone marrow cavities throughout the penguin body. Let's take closer look at these capillary beds. The structure to the left has interesting applications in penguin heat regulation. You can see oxygenated blood make its trip through the arteriole (on top, in red). At this point, blood can either enter metarterioles (running down along center) or through the smaller capillaries. Observe the small bulbs along the top arteriole at the beginning of each capillary. These are called pre-capillary sphincters. These are actually small bodies of muscle that control entry of blood into the capillaries. When a penguin isn't using certain tissues -- like when it is at rest or just walking around -- these sphincters contract and divert blood through the metarterioles. Occasionally, the sphincters relax and deliver blood to the tissues through the capillaries. But, let's say this is a capillary bed in the pectoralis major muscle, the primary muscle a penguin uses to flap it's flippers. And let's say the penguin is chasing after an elusive fish or trying to get away from a sea lion. That muscle needs a lot of oxygen. At this point, the pre-capillary sphincters relax and blood flows through the capillaries to perfuse the tissues with lots of blood and, hence, oxygen. When oxygen is released to the tissues, carbon dioxide leaves the tissues and enters the blood. Carbon dioxide then enters the red blood cells. With the delivery of oxygen to tissues, the loss of oxygen makes the blood turn purple in color (it's really not blue as the picture indicates). This is called the venous side of the capillaries. The venous capillaries empty into larger post-capillary venules and then into larger venules. Eventually these give rise to veins and return to the heart. At this point, the trip begins again, after being oxygenated in the lungs. Go onward to learn about penguin lungs.

The cardiovascular system, the system that delivers oxygen and nutrients to the cells of the penguin body, is the most important system in the penguin. Without a properly functioning cardiovascular system, a penguin can no longer think, mate, walk, swim or live. Therefore, nature has designed a pump and a system of blood vessels that are well-suited for the biological activities of penguins. Much like our own, this system is susceptible to deterioration. The cardiovascular system is a series circuit. Basically, this means that everything leaving the heart comes back to the heart sooner or later. Oxygen enters the blood stream in the lungs, travels by means of pulmonary veins to the Left atrium (a small chamber in the heart), into the left ventricle (a thicker pumping chamber in the heart), into the various arteries of the penguin and to the body. When cells extract nutrients and dump their waste products, blood travels in veins back to the heart in a similar fashion. So, what blood leaves the heart returns to the heart in due time.

There are several components to the penguin's cardiovascular system. The most prominent feature is the heart. The penguin heart, just like yours and mine, is a muscle. It's a well-designed muscle that contracts and expands the entire life of the penguin. The speed of the contractions increase when the penguin needs more oxygen (swimming, mating, fighting, and fleeing predators) and relaxes when the penguin needs less oxygen (resting, standing, sleeping). You can see in the image on the left a basic penguin heart. Penguin hearts and mammal hearts are unique among animal hearts. We all have four chambers in our hearts.

Basically, this means that oxygenated blood is always separated from poorly oxygenated (deoxygenated) blood. (This is considered an evolutionary advancement over reptiles, amphibians and fish.)These four chambers are the right and left atria and the right and left ventricles. The atria (singular: atrium) collect blood returning from the body or lungs and pump that blood into the ventricles. The ventricles have more important jobs; their either pump the blood throughout the penguin or to the lungs of the penguin so that it can be oxygenated. Since the left ventricle pumps blood throughout the penguin body, it is much stronger than the right ventricle which pumps blood to the lungs of the penguin only. It might sound strange, but penguin hearts need blood as well, just like all other organs in the penguin. A penguin's heart is composed of cardiac muscle and it has its own blood supply: the cardiac arteries. As in humans, these can become clogged and be the source of heart attacks and death in penguins.

From the base or the heart (opposite of the apex heart) arise the many arteries that supply blood to remote regions of the penguin and veins that drain blood from remote regions of the penguin. The common carotid arteries deliver blood to the neck, head and brain of the penguin. The jugular veins drain blood from those same regions after it has made its trip. Other important vessels arising from the heart are the axillary arteries, which deliver blood to the upper limbs (flippers) and the pectoral arteries, which deliver blood directly to the pectoral muscles of the penguin. We've already talked about three components of the cardiovascular system: the heart, the arteries, and the veins. Another component is the lymphatic system. Lymphatic vessels are small vessels like veins that collect fluid that builds up where it isn't supposed to be. Lymphatics deliver this fluid back to the heart so that it is returned into the regular (systemic) circulation. Although the lungs and respiratory system are intimately involved with the cardiovascular system, I'm going to give them separate treatment in the next section. A final component of the cardiovascular system is the renal system. The renal system is composed of the kidneys. While most people think of the kidneys as the means by which penguins excrete nitrogen and other wastes, the kidneys play crucial roles in maintaining blood volume, composition, pressure, pH (acid and base composition) and hormone levels. Without the kidneys, the penguin would die. Unlike human kidneys which are considered to have only one anatomical lobe, penguin kidneys have 3 anatomical lobes. An artery supplies each lobe.

The above circulatory map should give you an idea of the basic cardiovascular plan in the penguin. It is much more complex than this, and there are many smaller vessels that branch from these. (Furthermore, I've left out the veins.) However, you can see how all the important arteries course through the penguin. You're already familiar with the carotid arteries (observe how they enter the head and branch).

We've already discussed the axillary and pectoral arteries. A new artery is the systemic artery that arises from the systemic arch off the heart. It's very much like our aorta. It runs along the median aspect of the penguin and sends off several unpaired and paired arteries as it runs caudally. The first branch off the systemic artery is an unpaired artery: the celiac artery. The derivatives of this artery will supply much of the foregut of the penguin: gizzard, liver, pancreas, etc. There are also 3 paired arteries that deliver blood to each lobe of the each kidney in the penguin. After the renal arteries leave the systemic artery, femoral arteries are found. Femoral arteries supply blood to the legs of the penguin. Finally, posterior mesenteric vessels that supply the hindgut and caudal arteries supplying the tail branch. When tissues have extracted their nutrients, the blood will reverse its path in a similar fashion and return to the heart.

Penguins have lungs, too. But these lungs aren't like ours at all. You're probably familiar with our system of lungs: we have two and air is conducted in when we inhale and out when we exhale. These means that mammal lungs have blind ends. Air goes in, stops, and comes right back out. In penguin lungs, air never stops. Find out why below. Respiratory System Early in development, the respiratory system begins as a diverticulum, that is, an outpocketing, of the digestive tract. Shortly after budding outward, the digestive tract and the respiratory tract separate. The respiratory diverticulum forms two lung buds. At this point, human and penguin lung development are quite similar. Shortly after this point, penguin lung development becomes very different. Penguins develop additional organs called air sacs.

Penguins have to breathe air just like mammals; therefore, they have to come above water while swimming. Breathing begins with inspiration. Air enters the nares, the same thing as nostrils, or oral cavity (mouth). A short distance away from this site, air travels to the lungs through the trachea or wind pipe. Penguin tracheas are much like human tracheas. The penguin trachea is composed of mucous tissue, muscle, and cartilage, just like a human trachea. Here is what the penguin trachea looks like under the microscope:

The very top of the image is the lumen or tunnel portion of the trachea. Right below the lumen is the respiratory epithelium. It's amazing to think that the epithelium is only one cell thick! Beneath that is

some tissue and a thin layer of muscle. Dominating the image are two layers, or rings, of cartilage. Cartilage is tough tissue that sometimes forms bone. As air moves further down the trachea, it goes in two directions into the two lungs of the penguin. With this flow of air is a large volume of oxygen. In order to get oxygen into the blood, oxygen has to come within a very small distance of the penguin blood cells. This is when oxygenated air enters the parabronchus. The parabronchus is a specialized lung tissue found in penguins that isn't found in mammals. That's because air continually cycles through penguin lungs and air sacs. Oxygen passes along the cell surface of the parabronchus and diffuses into the tissues and then into the blood. At this point, carbon dioxide leaves the blood and enters the airspace of the parabronchus. Here's a picture of a penguin parabronchus:

This is how blood gets oxygenated in penguins. If you look at the last picture in this section, you can see a basic model of how air flows through the penguin respiratory system. It's similar, yet different, to our very own respiratory system.

You've just read about the cardiovascular system and how it distributes oxygen and nutrients throughout a penguin's body. The oxygen that the cardiovascular system carries has to be drawn from the atmosphere, and it is done so by the pulmonary or respiratory system. Unlike mammals, the penguin respiratory system has important functions other than respiration and ventilation. The principal differences between the mammalian pulmonary system and the penguin respiratory system is the distinct lack of a diaphragm, the continuous cycling of air, and the presence of air sacs. I'll discuss each of these individually and how they contribute to respiration. In mammals, the diaphragm is a muscle that separates the thoracic cavity from the abdominal cavity. Like all muscles, the diaphragm contracts. When the diaphragm contracts, it draws the lungs "open" which results in bringing in fresh air. When the diaphragm relaxes, the elastic properties of the lungs "close" the lungs and air is exhaled. This process is called ventilation. In penguins, respiration occurs with contraction of various abdominal muscles that compress the abdomen and relax the abdomen when contraction ceases. The figure above shows the complexities of the penguin pulmonary system. If you are familiar with the human lungs, you know there is a trachea, two mainstem bronchi and two lungs. That's basically it. Right away you can see that there are several other components to the penguin pulmonary system. While I trace the flow of air, start thinking about why this is efficient for birds like penguins who need continuous flows of oxygen. With the expansion of the abdomen, air is drawn into the trachea, which is continuous with the oral cavity via the oropharynx and the larynx. Air travels through the cartilaginous trachea. As it passes through the trachea, it runs through a distended portion known as the syrinx. The syrinx is the elaboration of the trachea that enables a penguin to make the numerous vocalizations that you hear like the ecstatic display or the trumpeting you might here when mates greet each other. The syrinx is also the site of the tracheal bifurcation, or site where the trachea splits into two channels. Each of these channels connects the syrinx to each lung. However, air first entering the respiratory system immediately flows into the posterior air sacs. All this occurs during the first inhalation. When the penguin exhales, air situated in the posterior air sacs flows into the lungs via the ventrobronchi and dorsobronchi. As air flows through these two bronchi, they travel out into smaller bronchi called parabronchi. You've read about this in the histology section, so you know this is where oxygen exchanges with the air and the blood. Unlike human lungs, the air doesn't stop here; it keeps flowing onward. When the penguin inhales for the second time, all the above starts over again. However, the air still in the lungs is pushed further along the respiratory system. Air in the lungs, parabronchi, ventrobronchi, and dorsobronchi is delivered to the anterior air sacs. At this point the penguin exhales one more time. The muscles of the abdomen contract and air is pushed throughout the system once more. Air sitting in the anterior air sacs is throttled into the interclavicular air sac, which is continuous with the trachea. Air flows out the trachea. The cycle of respiration is complete and it begins again.

So you can see from this sagittal image, respiration in penguins is truly circular. In effect, air is continuously flowing through the lungs with fresh oxygen. Instead of an in-and-out movement like that in humans, air flows constantly over the oxygen-blood exchange barrier. This proves to be a highly efficient means by which to extract oxygen. Penguins need this when they are "flying" through the water, evading predators or chasing after food.

An interesting fact to ponder: what would happen if you blocked the trachea of a penguin but exposed one of its upper limb bones to the open air? Although this sounds grotesque, there's an important anatomical fact in this experiment. The answer is that the penguin wouldn't suffocate. Why? Look at the second image on this page. Do you notice extensions of the interclavicular air sac running into each upper limb? Most major bones of the penguin, especially the humerus, are continuous with the air sacs of the body. Not only does this fill the bone cavities with air making them lighter, but it also serves as a reserve for air storage.

The last fact I will discuss regarding the penguin respiratory system is the role of the interclavicular and anterior air sacs in thermoregulation.

You've read throughout the species pages that penguins consume various marine animals, such as small fish and crustaceans, as food sources. This process of "foraging" is energetically expensive and can be dangerous. However, when they do feed, the food must be digested. Therefore, it travels through the penguin digestive system, which is a lot like the digestive system of all birds. It contains 6 primary components: esophagus, crop, stomach, gizzard, intestine, and cloaca. Digestive System All penguins form from a single cell, the zygote. Shortly into the development of the penguin, the embryo folds over itself and forms a continuous canal with a single opening and two close-off openings. The single opening leads directly to the yolk sac, and the other two closed openings will form the oral cavity and the cloaca. This is the formation of the alimentary canal, or digestive tract. Formation of this structure is better explained in the embryology section. The oral cavity of the penguin contains a few structures that you are probably familiar with. It is covered by the bill or beak, which is a hard, keratinous structure. Penguin bills vary in shape and size, and this is dependent on the genetics of the penguin and how the penguin has evolved. Internal to the penguin bill is the oral cavity proper. Structures found within the oral cavity proper include the tongue with interesting keratinous protrusions emanating from it, and a cleft palate (roof of oral cavity). These keratinous spikes protruding into the back of the oral cavity help penguins hold on to fish or crustaceans that they capture when fishing for food. The cellular lining of these structures is stratified squamous epithelium, which are basically layers of flat cells that are designed to resist abrasion. The oral cavity leads into the esophagus, a structure which conducts food to the crop and eventually the stomach. The crop can store food for penguin chicks or for when the penguin wants a snack later on. Food partially digested in the stomach is delivered to the gizzard, a thick, muscular structure which grinds food into smaller bits. After this, the digested food is shuttled into the intestine to be absorbed into the body. The intestine of the penguin is similar to the intestine of the human being. The lining of the intestine is covered by numerous structures called villi. The picture below elucidates the basic villus. A villus is a projection of cells that increases the absorptive area of the intestine, and the cells covering the villus are renewed about once a week. Inside the villus are blood vessels and lymph vessels that shuttle digested nutrients into the body.

Enterocytes, or absorptive cells, take up sugars, proteins, minerals and liquids from the mix of digested fish or crustaceans penguins eat. After this occurs, sugars, proteins and minerals are conveyed into the capillaries and lacteals of the villus. Digested food particles enter the blood stream and are conveyed to the cells throughout the penguin. Here is a real picture of a penguin intestine (specifically ileum). You can see the villi on the right of the image and the muscular layers on the left of the image.

Food is digested and absorbed all along the intestine. The fate of undigested food or undigestible food is to end up excreted in the feces. Undigested food substance is deposited into the cloaca. A cloaca is a common dump found in penguins (and in embryological humans, even). Since this is a common dump, the excretions of the kidneys (nitrogenous waste in the form of uric acid) are deposited here. Additionally, when mating, sperm leaves the male's cloaca and enters the female cloaca. So, as you can imagine, the cloaca has a lot of functions.

In order to survive, a penguin must consume food to extract the vitamins and nutrients from that food. Vitamins and nutrients are necessary components of penguin metabolism; penguins uses these components for the synthesis of proteins, DNA, and RNA. Other components are essential intermediates in metabolic reactions. They give penguins the colors that you see and enable penguins to perform the daily functions that are necessary for survival. Like the cardiovascular and pulmonary systems, the gastrointestinal system of the penguin is catered to bodily needs. There are several specializations that enable penguins to store food for chicks; there are specializations for pulverizing food since penguins have no teeth. Before undigested food exits the penguin, it follows a twisting path to ensure that energy extraction is maximized. Look at the diagram below. Note the components. I'll talk about them individually. Food first enters the oral cavity. If you've visited the histology section, you know this is the "mouth" of the penguin and it is the initial site where food begins its digestion. However, very little digestion occurs in the oral cavity -- probably none. There are, however, mucous glands that secrete mucus, aiding the passage of food. The oral cavity assumes a role of connecting the outside world to the esophagus, which is a conduit from the oral cavity to the crop. The esophagus is a tube-like structure that has a muscular wall. Through movements known as peristalsis, wave-like contractions propel food from the oral cavity into the crop where food sits if it is to be regurgitated or if the stomach is full. It is my understanding that the crop is merely a diverticulation (distended pocket) of the esophagus and may not have any specialized functions, per se. Food leaving the crop either reverses the esophagus and out the oral cavity or it passes through the proventriculus. The proventriculus is an important component of the digestive system of the penguin since it secretes digestive enzymes that allow digestion of fish and crustaceans to begin. The proventriculus is the first part of the penguin "stomach." The second part of the penguin "stomach" is the gizzard. You all may be familiar with the gizzard's function by now; it grinds and pulverizes fish and crustaceans into smaller components that are more easily digested by penguins. The gizzard has a thick muscular wall (this is necessary to produce the contractions which grind up food). The internal surface of the gizzard is similar to sandpaper since it contains a grainy keratinous substance known as koilin. Histologically, this is known as the cutica gastrica. Once partially digested food exits the gizzard it encounters secretions of the liver and the pancreas. The liver, located inferolateral to the heart, produces and secretes bile, a collection of acids, pigments and cholesterol that are vital to the digestion and absorption of fat. To curb the acidity, the pancreas will secrete sodium bicarbonate to neutralize acid. In addition to this buffering material, the pancreas will also secrete various protein-digesting chemicals, sugar-digesting chemicals and fat-digesting chemicals. This all occurs once partially digested food enters the intestines. Therefore, as you can imagine, most digestion and absorption occurs in this region of the gastrointestinal system. Food travels along this curvy organ while it is being slowly digested and absorbed. When material can no longer be absorbed, it is delivered to the cloaca of the penguin. Here the feces meets up with uric acid secreted by the kidneys. The contents are expelled at the convenience of the penguin.

We left our study of penguin histology with a close look at bone. Bone forms the fixating points for skeletal muscle (or most skeletal muscle anyway). To generate the powerful movements require to whip through the water, penguins recruit muscle fibers to propel themselves. Here's a closer look... Muscle Skeletal muscle is invested by a thin connective tissue blanket composed of collagen and elastic fibers. Remember, collagen was the same fiber used to form bone. Collagen is particularly strong and is well suited to be part of the musculature. Muscle is composed of the following components: Epimysium Perimysium Fascicles Endomysium Skeletal muscle fiber Sarcomere cross-bridges

Don't let these terms scare you off, for they are simple concepts once you look at a histological section of muscle. On the right, you can see a cross-section of penguin muscle. The light pink and tan colored bodies you see distributed throughout the image are actual muscle cells, also known as muscle fibers. If you look closely, you can visualize the eccentric nucleus of each muscle fiber. Look for the small purple dots on the sides of each muscle fiber. Running between these tan and pink bodies is the endomysium, a thin layer of collagen tissue that separates each cell from one another. If you look in the upper right, just off the center of the image, you can see a large empty space of collagen tissue. This is part of the perimysium. Perimysium is thicker collagen tissue that creates fascicles, or bundles of muscle fibers. Several fascicles rolled up together create the muscle that you can see and feel. These bundles of fascicles are surrounded by what? If you guessed epimysium, you are correct. From the outside-in, it goes epimysium to perimysium to endomysium. Penguin skeletal muscles are under voluntary control. That means that they won't function unless the penguin wants them to function. In order to get muscles contracting, the brain sends signals through motor nerves. Nerves consist of many, many neurons, or single nervous cells. As a neuron descends from the central nervous system, it branches (i.e., ramifies). Each branching of a neuron connects with each muscle fiber. One motor neuron services one motor unit, a collection of muscle fibers. When stronger contractions are needed, e.g. when a penguin needs to escape a predator, more motor units are recruited.

As more motor units are recruited, more muscle fibers contract and create stronger muscle contractions, propelling penguins. The image on the left was performed with a scanning electron microscope. You can see a "thick" muscle fiber descending from the upper left down to the lower right. A thin wisp of a neuron (N) can be seen winding down from the upper left and synapsing with the muscle fiber at the myoneural junction (MJ). There are many muscles in penguins that are involved in swimming, and you will learn more about them in the anatomy section. The basic operation for all skeletal muscles is the same. After a synapse occurs, a signal is given to each muscle fiber to contract. Within each muscle fiber is a bundle of filaments. These filaments are made of proteins called actin and myosin. The two of these muscle fibers make up the sarcomere. The sarcomere is the functional unit of the muscle.. The sarcomere shortens each time the muscle contracts. When the actin and myosin pull on each other, the muscle shortens. When the muscle shortens, this is called contraction.

There are varying forms of aggression, but first I would like to point out the difference between displays of aggression and actual aggression. Both the displays and acts of aggression are encompassed by the term behaviour; however, displays tend to be signals that have evolved, and those displays that are maintained may actually communicate something, such as an impending act of aggression. This individual may or may not act upon receiving those signals. Further, these signals may be tools used by animals to assess the strength and "will" of potential competitors.

For a number of years, this behaviour was hard to assess. As the image below suggests, the acting penguin (either a male or female, generally at the nest) tucks its head under one wing and raises the opposite wing in motion. The penguin may vary this behaviour by growling or by moving its head to the opposite side of the body. Although males and females perform the billto-axilla, it is given most often by males and it seems to indicate ownership of a general area.

However, this behaviour does not seem to be directed towards particular individuals, rather groups, neighbors or passer-bys.

There are two types of stares: the sideways stare and the alternate stare, the latter being the more intense and aggressive behaviour. Penguins have poor binocular vision, and some penguins have no binocular vision at all. For this reason, the sideways (and alternate) stare act as methods for focusing on the threatening object(s). These stares may be performed while lying down or while standing. In either case, the head of the staring penguin is placed in such a position that affords it ample time to strike with its bill. The flippers are kept close to the body but (as I have noticed in the Humboldts) may extend to either preserve balance or appear more threatening while leaning forward. These stares are given by territorial penguins. The sideways stare may also be given by females as they approach the nests of unpaired males who have been giving Ecstatic Displays (see Displays Between Sexes).

The alternate stare serves the same purpose of the sideways stare but is separated from the former because it is employed during more intense situations. The focal penguin may perform an alternate stare if an intruder persists in the territory or if neighbors tend to move about frequently near the nest.


The point and gape are closely related and often the gape is the logical progression from the point behaviour. Both these behaviours are more aggressive than the stares or bill-to-axilla. The point is a simple behaviour recognized by Penny (1968) which involves, as the name implies, the penguin's pointing (with bill) directly at the threatening object. Generally given in the pointer's territory, the object of the point may be intruding or moving very closely to the territory. Often the object of the Point is another species. The Point does not have to be given within a territory and may be employed to warn others when they approach too closely to the pointing individual. "The Gape" was originally seen as part of the Direct Stare that Penny had observed but is widely regarded now as separate, though related, behaviour. The gaping penguin often employs the same offensive (or defensive) posture as it does while pointing, opens its beak and emits a loud, growling sound. This gaping may culminate in what is known as Beaking. Gaping individuals may rock back and forth until they both lock beaks and begin a bout of tug-of-war.

The Charge is seen as the most reliable indicator of attack, since it involves the most intention. As the name implies, the focal bird "charges" the threatening object, though it may not attack. Generally, the penguin leans forward, extending its wings and opening its bill, while it locomotes toward the other penguin. Penny (1968) noted that this movement may be silent or may involve general squawking. Penguins incubating eggs or guarding chicks will not, for obvious reasons, leave the nest to charge an approaching individual.

Displays between sexes refer to the acts involved in mate acquisition and pair-bond maintenence. Some refer to this activity as courtship or "ante-nuptial" behaviour, and much of it is. However, some of these displays extend well beyond the social bond. Richdale called this activity "love habits," but this seems to connote too much emotion. And the word "habit" suggests that the activity is fixed and, as a term, "fixed action pattern" does not adequately describe these behaviours.

By far the most dramatic display between the sexes is the Ecstatic display. The name was coined by Wilson (1907) but as it applies to behaviour, the term has been somewhat modified over the last century.

The Ecstatic display is given typically by unpaired males attracting females. The calling may be unprovoked or incited by the approach of other penguins (and humans!) or may be a consequence of mass ecstatic displaying. The focal bird aligns his body vertically and pumps his chest several times. After this motion is complete, the individual emits a harsh, loud breying sound while arching his flippers back as far as possible. Quite audible from far distances, the ecstatic display is likened to the lone trumpet call (see Trumpet). There are several variations on this theme, the most common being a side to side movement of the head while calling. The call may also be used by individuals guarding chicks or by paired males. The Ecstatic display is also divided into two sets of varying intensity: the intention ecstatic and the imploring ecstatic. Both are similar in function to the ecstatic display, yet the former is incomplete and the latter is an amplified ecstatic display.

The Bow, like most other forms of behaviour, was first named and described as an Adelie penguin behaviour. The Bow is a behaviour demonstrated most often in pairs at the nest. In a normal situation, the female will bow to the male after approaching the nest. Conversely, the male may do the same. Penguins may also walk about the area while bowing. And thus, Bowing is considered to be both a form of appeasement and a tool for maintaining social bonds.

Divided into Loud Mutual displays and Quiet Mutual displays, these displays are seen shortly after one of a pair has returned to the nest after a time of absence. The focal penguin approaches the nest and calls out to its mate. The mate then approaches and leans toward its partner and calls in time; following this, the two stretch in the vertical direction to end the calling. One of the pair may shake its head during this event. For the quiet mutual display, much of the behaviour is the same, save the noise. One issue may separate the two - a loud mutual display may be given by a lone penguin.

In my own study I often find this hard to distinguish this from the mutual display. By comparing what Warham has to say about the trumpeting performed in the crested penguins (Eudyptes) with how Spurr describes the mutual display in Adelie penguins, I have concluded (at least for the time being) that mutual trumpeting differs in that the members of the pair stand side-by-side and may lean forward and then upwards while performing the behaviour. It is my understanding that the mutual trumpet and the mutual display serve relatively the same functions: mutual recognition and greeting after long absence.

The vertical head sway is yet another general behaviour displayed mostly by males to indicate ownership of a nesting territory and possibly to display for females. The behaviour is initiated when a penguin bows his head toward his feet and rapidly raises it back to the vertical while emitting a growl. Once at the peak, the penguin sways his head from side to side; during this motion, sounds may be heard from the penguin. The vertical head sway, like most penguin behaviours, varies in intensity between penguins. Certain species greatly exaggerate the extent to which the head is swayed, while others the speed. I have noticed that the Humboldts (if indeed their behaviour is a "Vertical Head Sway") rock their heads in a rapid, albeit jerky, motion encompassing full 180 degrees of motion from the vertical.

Two forms of preening fall under the category of behaviours demonstrated between sexes: allopreening and mutual allo-preening. The former involves one individual preening the other. The act may be a form of recognition, reconciliation or a behaviour used between individuals of a pair. When the recipient penguin also preens the focal penguin, this act is known as mutual allo-preening. This event is sometimes known as the "kiss preen" since the focal penguin usually preens around the neck and beak of the recipient penguin. Most often this is done between the members of a pair and is a form of recognition. Of course, the goal of most sexual displays is to gain mates or to reinforce the pair bond. What proceeds is a brief summary of reproduction in penguins. Although I will present drawings that elucidate

the sexual organs, I'm not going to develop them thoroughly. Besides, I don't know much about the specific physical mechanisms there. Like most birds, the reproductive cycle begins with the courting displays. You've seen those types of displays above. Now let's get into the thick of it.


The reproductive structures in penguins do not differ extensively from those of other avian families. All birds reproduce sexually: males transfer sperm to the female through the cloaca. During copulation, the event of contact between the two cloacas is known as the cloacal kiss. Reproductive structures of the male penguin The following discussion focuses on general avian reproductive structures, not penguin reproductive structures specifically. Although penguins differ quite a bit from other birds with respect to taxonomy, they are birds, and it is not unreasonable to suggest that their reproductive organs are similar. There are four principal parts to the male reproductive tract: the testes, the epididymis, the ductus deferens and a sperm storage location near the cloaca (the common orifice). The testes are located dorsally.

Only in a few species of birds does the male possess an intromittent organ; these birds include ratites, kiwis, cracids, screamers and waterfowl. Penguins do not have intromittent organs. There are varying reasons as to why intromittent organs would be evolutionarily advantageous in these species, but they will not be discussed here. Stage 1 (n=16 cases observed). A row of spermatogonia and Sertoli cells ring the seminiferous tubule next to the basement membrane. Primary spermatocytes are present in an irregular row toward the lumen. The lumen is filled with sytoplasm. Stage 2 (n=20). There is a basal row of spermatogonia and Sertoli cells. Up to one-half the primary spermatocytes in a given tubule cross section are in the stage of synapsis where chromatin is concentrated to one side of the nucleus. Stage 3 (n=2). A basal row of spermatogonia and Sertoli cells ring the basement membrane. Towards the lumen two to three rows of primary spermatocytes are in synapsis. Stage 4 (n=2). A row of secondary spermatocytes lies toward the lumina of tubules. Peripheral to these are cells of previous stages. Stage 5 (n=7). Spermatids are present surrounding the lumina. Intertubular cells are almost completely confined to the triangles formed by the meeting of three tubules. Stage 6 (n=10). Rining the lumina are bundles of spermatozoa with their heads pointing toward the basement membrane. The lumina are filled with cytoplasm. This stage represents breeding condition.

Stage 7 (n=10). In this, the climactic stage of spermatogenesis, spermatozoa and cytoplasmic particles are present in the lumina. Stage 8 (n=13). The stage of degeneration where the various cells from all stages in spermatogenesis, except spermatogonia, discharge into the lumina.


In most birds, the female has one ovary. This ovary produces ova that over the period of receptivity increase to 1000 times their original sizes. The basic female reproductive tract includes the ovary, the magnum, uterus, uterovaginal junction and the oviduct (of the cloaca). Shortly before the period of egg laying, the ovary carries several developing follicles, each known as the ovum. Since each ovum develops within 24 hours of the pervious, as "hierarchy" of follicles is created and the larger the follicle, the sooner it is released. This is the crucial phase of receptivity in birds, and though it may vary from species to species, it is 15-30 minutes in length (Warren & Scott, 1935). Following this stage, the perhaps egg passes through the magnum, the isthmus and through the uterovaginal junction while collecting albumen and becoming calcified.


In the Adelie penguin, birds return to the rookeries in the late "autumn" and acquire pairs. Though being early does not confer an assured mating, the early arrivals are usually the ones who do achieve mating status. During this time, Adelie body weight is high and over the next month or so of breeding, they can lose up to 1/3 of that weight. After the egg laying period has ceased, the female leaves the rookery for the sea while the male stays to incubate the egg; during this time, males remain faithful to their fatherhood duties and do not consume any food. Because this devotion is so extensive, succeessful breeders are the penguins who are excellent foragers and can amass sufficient fat reserves for this six week period. D.G. Ainley's data for the breeding Adelie show that young birds do not breed (2 years of age). Females may begin to breed at age 3, but males do not begin to be successful until age 4. By age 6, most of the females in Ainley's experiment had bred at least once but it was not until age 8 did the

males breed at least once. At this point, I should point out that there is a difference between being physiologically capable of breeding and being socially capable of breeding. Young birds may have the mechanics required to produce offspring, but their relative inexperience with acquiring mates and caring for the young makes them unsuccessful breeders. The process of acquiring a mate may be extensive and require several displays before choice occurs. In the Adelie penguin, there are some four different displays observed in acquiring mates and maintaining the social bond: bowing, mutual displays, ecstatic displays and allo-preeing. Ultimately, these acts lead to coition (cloacal contact). The process involves mounting and finally copulation with the female. The mechanics are different and vary among penguin species, but that for the Adelie is explained here. Coition or copulation is not required for mounting to be successful. Mounting simply refers to the act of the male standing on the female's back and attempting a copulation. This mounting behaviour is typically preceded by the arms act. During this event, the male approaches the female from behind and beats his wings on her sides. During this motion, the male may also rub the underside of his head and neck on her back and on the top of her back. As the male continues to vibrate his wings on the female, the female typically turns her head so that their bills vibrate next to or on each other. This act is followed by the male's lowering of his tail; the female in turn raises her. If the mount and copulation is successful, the two cloacas will make contact and sperm will be transferred. In Adelies, this act will last for a minute or more. In the Humboldt Penguins I have observed, the arms act and copulation is very similar; however, I have not seen the female to rotate on to her back as this picture suggests. Mounting and copulation typically takes less than 3 minutes in my Humboldt group.
Spurr, Dr. E.B., Communication in the Adelie Penguin in Stonehouse, The Biology of Penguins

If you've read the section regarding sexual behaviour, then you read a little bit about the reproduction of penguins. In this section, I'll give it serious development. Penguins reproduce by sexual means. "Sexual reproduction" implies that two gametes -- single cells -unite to form one single cell or zygote. The female gamete is the ovum and the male gamete is the sperm. After copulation occurs and assuming sperm make it to the magnum of the female reproductive tract, the sperm penetrates the ova. This is called fertilization. The resulting, unified cell is called the zygote. Look at the drawing to the right. First we'll trace development and the path of the sperm. Sperm develop in the testes; penguin males have two of them closely apposed cranioventral to the first kidney lobe in the abdomen. (Remember the terms cranial and ventral?) The sperm then travels into the ductus deferens which runs in medially and enters the caudal cloaca (bottom of the cloaca) in a specific region known as the urodeum. During copulation, cloacal contents are delivered to the female cloaca. If the copulation is timed with ovulation, fertilization may occur. Females usually have only one ovary; usually the left ovary persists. A developing follicle supports a developing oocyte (immature ovum). As the oocyte develops, fluid built up into the follicle raises intrafollicular pressure to the point of rupture. The oocyte is expelled into the body cavity. Ovulation occurs, releasing an ovum into the body cavity (technically this is called the peritoneal cavity, but we won't go into that distinction). The ovum is swept into the infundibulum by the fimbria of the infundibulum. Fimbria are finger-like projections of the infundibulum that become engorged with blood during ovulation. The maturing ovum moves into the magnum where it encounters the sperm. Fertilization occurs and the zygote, or "egg," continues to travel down the female reproductive tract. As the egg continues its travels, it acquires a layer of albumen and the shell membrane begins to deposit in the isthmus of the tract. All the while, the size of the yolk sac and egg is increasing. As the egg passes through the uterovaginal junction, it hardens extensively and is passed into the world through the cloaca. The picture to the left is of an early embryo. Quick inspection will show you that the embryo at this stage is overwhelmed by the extensive yolk sac which in this case imparts a yellow color to the entire contents of the egg. It is from the yolk sac that the developing penguin embryo draws all its energy. There's plenty of necessary fats, cholesterol and sugars in this area of the egg. Developing neurological systems require fats and cholesterol, while most energy is derived from the metabolism of simple sugars like glucose, galactose, and fructose. You can see an extensive network of blood vessels stemming from the embryo to the recesses of the yolk sac. Nutrients enter the blood vessels and they traverse to the embryo proper. At this point, nutrients can be distributed throughout the developing embryo.

Within a few days, the embryo grows in size to resemble something like a animal form. The embryo to the right has been incubating for several days. At this point, it is still drawing nutrients from the yolk sac. Its oxygen is supplied from the external world; it enters the egg through microscopic pores in the shell. Metabolic carbon dioxide exits the shell by these same channel pores. The picture to the right shows the developing embryo. The most prominent feature of this embryo is the developing eye. Close inspection will reveal unrotated limb buds (during development, upper limb buds will rotate laterally and lower limb buds will rotate medially). Vascularization (blood vessel development/presence/etc.) is extensive during this stage. The primary artery delivering nutrients to the embryo can be seen in the median plane of the embryo, entering the ventral surface. The dorsal surface of the embryo is unremarkable in this picture, but the vertebral column is developing.

A slightly earlier staged embryo. Several features are prominent in this sagittal section. Moving cranially to caudally, we can see the head of the penguin developing its eye and brain (forebrain). At this stage, the heart is prominent as a ventral diverticulum of the ventral surface. At about this level on the dorsal side of the penguin embryo is the developing auditory center. It is indicated here as the otic vesicle. Running through the dorsal surface of the penguin embryo from cranial to caudal regions is the neural tube. The neural tube develops from the ectoderm of the embryo. The ectoderm is the outermost cell layer of the embryo. The neural tube formation is induced by the notochord, a structure shared by all Chordate animals. (Not all Chordates develop vertebrae, however. That is why birds are listed with the sub-phylum Vertebrata to note that they are Chordates with vertebrae.) The neural tube will develop into the central nervous system which includes the spinal cord and the brain. The notochord will regress into the intervertebral discs. The last feature of this embryo section are the somites. Somite function can be difficult to understand. They are condensations of cells around the neural tube that form the vertebral system. They unite with the notochord and surround the developing spinal cord. Once they coalesce around the spinal cord, they fuse together to form the vertebrae and the base of the skull. The development of the vertebral column and base of skull is actually more complicated than

that, as they have to permit exit of nerves from the spinal cord throughout the length of the spinal cord, so the somites don't completely fuse around the spinal cord. Incubation period varies among penguin species, but the chick eventually hatches from the egg if the conditions are right. I won't go into chick development, but know that there is a gradual process of growing and plumage changes before the chick becomes a juvenile and before the juvenile becomes an adult. Each chick goes through the following stages: natal down to juvenal plumage via the prejuvenal molt. From the juvenal plumage, the penguin moves to its basic plumage via a prebasic molt.

1. What is a penguin? A penguin is any bird that belongs to the Order Sphenisciformes. That would be a strict definition. But, that doesn't mean much to an individual upon inspection. First of all, a bird is any animal that is a homeothermic vertebrate animal with a pair of modified upper limbs that support or supported flight at one time in the evolutionary development of the animal. A penguin would be any animal that belongs to the class Aves, which contains all birds, is incapable of flight due to loss of the proper machinery, is naturally restricted to the Southern Hemisphere, moves about by swiming and walking, nests on land, and shares similar morphological features -- mainly color and striping patterns -- between the sexes. 2. Where does the word "penguin" come from? The origin of "penguin" is nebulous. Sources suggest that it comes from the Swedish alka from which the English "auk" was derived. From this term came the expression "great auk" or "penguin." However, both "great auk" and "penguin" did not describe what we know as penguins, rather great auks, another type of flightless bird (which happens to be extinct). It is also theorized that "penguin" comes from the Welsh expression pen gwyn meaning "white head" or from the Latin pinguis, meaning "pin wing." Both these terms seem unlikely origins for the word "penguin," however. I don't think anyone knows. 3. How many species of penguins are there today? The number will vary depending upon whom you ask. Some say at least 18; most say at least 17, while others admit to only 16 or as few as 13. I think there are 16, or at most 17, species. The issue of speciation is debated on whether certain populations are merely hybrids of two species or distinct species. Speciation is said to have occured when two populations of animals cease to breed and develop on their own evolutionary paths. What is the question here is whether certain "species" of penguins are still capable of interbreeding. Two issues of contention are the Little blue penguin and White-flippered penguin whose morphologies are so similar that many will argue that they are conspecifics with subspecies differentiation. Others contest that the Fiordland and the Snares Island penguins are the same species and should be arranged into subspecies. Some say the Royal penguin is a geographically distinct race of the

Macaroni penguin. Still others contest that the extent of differences in the genus Spheniscus does not merit their separation into four species (this is not a popular view). 4. Then how many species of penguin are extinct? From the Late Eocene: 7 known species. From the Early Oligocene: 7 known species. From the Early Miocene: 7 known species. From the Late Pliocene: 2 known species. More development of this is dealt with in the section entitled Natural Evolution. Please visit this section for more complete coverage of penguin evolution. Are there any recent extinctions? No, but here is a list of the status of current penguins:

5. Why are penguins the only birds in the Order Sphenisciformes? Some biologists contest this designation of taxonomy. Others argue that penguins are so unlike other avian species that this arrangement is warranted. The biology and evolution of penguins is such that no other bird amongst the neornithes even closely resembles them. Because the penguin is unlikely to be relatives of these birds except distantly, scientists have placed them into their own order. 6. I noticed that the root "sphenis-" is in the taxonomy of the order, family and in one genus. Why? Spheniscus demerus, or the Blackfooted penguin (also known as the African or jackass penguin) was the first penguin discovered by European explorers. Hence, the root "sphenis-" was applied.

7. Where do penguins come from? Basically, there is little evidence that penguins moved from any particular place to their current locations. Early fossil records date the arrival of penguins in the late Eocene - 37 million to 45 million years ago. Fossils of this period were found in Australia, New Zealand and Antarctica, all current locations of several modern day penguins. Some of these fossils suggest penguins were as tall as 6 feet! So, it is likely that a distant ancestor of penguins migrated, either by flight or by swimming, to the southern hemisphere. This common link between modern penguins and ancient flying relatives has not be found; only penguin-like fossils have been found in the southern hemisphere. More on this in the section on Natural Evolution. 8. How closely are penguins related to other birds? It depends on which birds you are talking about. To remind you, Neornithes developed into Paelognathes (old jaw) and into Neognathes (new jaw). From this group of birds developed some other specifics, but the two of interest to us are the Chardriiformes and the Ciconiiformes. Notice that from the Ciconiiformes stem the Pelecaniformes (obviously containing pelicans), and from the Pelecaniformes stem the Procellariformes, which gave rise to Graviidae (loons, etc.) and Spheniscidae. There's Spheniscidae, our old friend the penguin. So, you can see that the auks, which are often mistaken as being closely related to the penguin due to their similar size, shape and ecological niche, aren't closely related to penguins at all. However, you can see what are closely related to penguins: the Procellariformes from which penguins descend directly, and the Graviidae, which is a daughter clade of the Procellariformes with the Spheniscidae.

So, you can see how different, based on the phylogeny chart, penguins are from modern birds. Despite the fact that they don't fly either, ratites (ostriches, emus, rheas) aren't closely related to penguins. The closest relatives of penguins are albatrosses, petrels, and shearwaters. They are all Neornithes. It seems ironic that even though they look like penguins, puffins, razorbills and murres aren't related to penguins either. 9. What is the full scientific name of the _____ penguin? It depends on which birds you are talking about, but a generally complete name would include the major phylogenic classifications. For example, the Humboldt penguin would be called (technically) Animalia Chordata Vertebrata Aves Sphenisciformes Spheniscidae Spheniscus humboldti. Notice that all the names except the species name (humboldti) are capitalized. The species names are not capitalized. Furthermore, because they are Latin terms being used in English text, they should always be italicized. You would abbreviate the Humboldt penguin's name with the following: S. humboldti. (Just like you would abbreviate Tyrannosaurus rex with T. rex, not T-Rex, not T-rex, not T. Rex or any other. 10. What is the name for a group of penguins? I don't think there's a fancy name like "gaggle" (geese) or "rafter" (turkeys). The name for a group of penguins is simply "colony." 11. How do penguins survive in such cold temperatures? Ironically, some penguins actually overheat in these regions. It is a noticeable trend that the colder the region, the larger the penguin. This trend is based upon a simple physical property that the smaller the surface area to volume ratio, the smaller the extent of heat loss. In other words, if you have a lot of surface area, you have a lot of area to interface with the cold air and cold water. Therefore, it is easier to lose heat. Little-blue penguins have the largest surface area to volume ratios, so they must live in warmer climates. Emperor penguins have the smallest surface area to volume, and they retain heat easier than the smaller penguins. However, emperor penguins do have problems retaining heat. That is why they form creches (huddles) during the winter. I'll develop more on creches in the section on Deconstructing Penguin Myths. Creches aren't necessarily what you think they may be. In addition to body size, penguins in cold regions amass fat under the skin which performs as an excellent insulator. Nonethless, this layer is relatively thin when one compares it to seal or whale blubber in the whole body ratio. However, it is of primary importance that all penguins maintain their feathers. Feather integrity is crucial to heat retention, and it is the primary superficial (external) mechanism whereby penguins regulate heat. Feathers "pocket" air between the skin and the environment, much like insulation in your house. The fiber glass or foam insulation has many airspaces which trap air, and in the case of penguins, this air is warm. Lastly, birds are homeotherms, just like us. This means that their heat is maintained by their metabolism. They generate enough heat (normally) to keep warm and the fine regulation of metabolism of their bodies insures that their heat remains in a constant range. However, penguins have been noted to actually drop their body temperatures when entering the cold water. This biological event minimizes the heat loss of the body. 12. Why are penguins flightless? Other than the obvious fact that their weight to wing area ratio is incapable of supporting flight, it is theorized that nature selected penguins to be flightless. Theory suggests that in penguins (and other

flightless birds) food was readily accessible and that the long search flights for food were not required. Consequently, nature did not select for wings capable of producing flight. This assumption, of course, is based on the idea that all birds originated from an ancestor that flew. There is little evidence that suggests that penguins followed some alternate lineage. More on this is treated in the section on Natural Evolution But as many people have observed, penguins do "fly." Penguins use their wings, more appropriately called flippers, to propel themselves through the water; to steer, penguins use their feet as rudders. 13. If penguins swim, their bodies must be well adapted to aquatic life. Is this true? Yes, they are tremendous swimmers, but contrary to popular opinion, penguins do not achieve relatively high speeds within the water. Stonehouse suggests that speeds range anywhere from 10-12 mph. Likewise, the diving capacity of penguins seems to be overestimated as well. The record dive for a penguin was recorded at 1772 feet for 18 minutes. This feat was performed by a rather "talented" emperor penguin and is not representative of stock. Penguins typically exhale before submerging and only stay under for a few minutes, barely exceeding the time the average human being can submerge himself. Additionally, penguins dive to depths of less than 60 feet. 14. Do penguins have teeth? Why not? No, penguins don't have teeth. In evolutionary past, birds did have teeth, but over time they lost them to reduce weight. The more an bird reduces it's weight, the more able it is to fly. But, you may be thinking right now: penguins don't fly! That's right; they don't. However, penguins evolved from birds that did fly. Teeth are also adaptations for grinding food. Penguins, and all other birds, have gizzards which are thick, muscular structures that grind food in place of teeth. 15. Is a penguin beak made out of bone? If you saw the picture of the penguin skull in the histology section, you might have thought that penguin beaks are made out of bone. This is not the case. Penguin beaks are made out of keratin, a protein that consists of two braided strands of smaller proteins. It's the same protein that makes up penguin scales, human finger nails and human hair. Penguin feathers are also made out of keratin.

16. Why don't penguins get sick from drinking sea water? Penguins don't drink Poland Spring or Evian every day, and some of them don't have freshwater sources since they are marine animals. The seas and oceans are high in salt concentration. If we drank salt water, we'd become very sick. Penguins don't get sick. Both penguins and other marine birds have salt glands located beneathe their skulls. These nasal glands have many little glands inside them that are surrounded by capillaries. As blood flows through these nasal glands, salt leaves the blood with just a little water. This high salt water drips through ducts that exit through the nares (nostrils). Often you can see penguins shaking their heads with what looks like gooey drippings flying from their noses! This is actually salt with water. See how the gland works below:

17. Do penguins get sick? Absolutely. They are biological systems just like we are and are susceptible to bacteria and viral infections, as well as the normal maladies that affect some of us: deteriorating eye sight, cardiovascular disease, etc. However, two of the more common infections penguins get -- and this is regionally dependent -- are infection by Aspergillus fumigatus, a fungus, and Avian malaria parasite. Spores of the Aspergillus fumigatus fungus typically causes lung infections, which are demonstrated by shortness of breath in penguins. A. fumigatus typically hide in fecal matter and moist, anaerobic conditions. When they infect penguins, the penguin usually becomes malnurished (if it isn't already) and dies. Immunocompromised penguins die sooner than others. Avian malaria is another common infection in the Malaria belt (Africa), but it more common affects captive species. Aspergillosus and avain malaria are both much more common in captivity than in the wild. Most indications are that wild penguins die of predation or the common conditions that kill penguins "naturally." 18. Do penguins have parasites? It depends on the species. Species like the Adelies and Emperors generally don't have to worry about this sort of thing since their climate is ultimately icy and cold. Species like the Rockhopper or Gentoos that live in the Periantarctic ring generally don't have to either. However, those penguins that nest in forests or in heavy grassy areas probably have problems with them. There are two particularly pesty parasites that infest penguin nests and penguins in general. The first is the Mallophagan louse. This louse is of the breeds of sucking lice. Generally they subsist on penguin blood and other exudate that leaves the penguin via the penetrations the louse creates. Lice can also lay their eggs in the skin or feathers of the penguin. Another parasite is the Itch mite. These parasites are about 0.5 mm in width and are found in penguin feathers. Small amounts of either of these parasites generally aren't a problem; however, heavy infestations can have serious health consequences for the penguin.

19. I saw a penguin at the Zoo. It let out a sound like the braying of a donkey. What was that? You may have seen any memeber of Spheniscus performing a courtship display. Many penguins perform what is known as an Ecstatic Display, a display in which the penguin arches backward, throws back his wings, and brays like a donkey into the air. Ecstatic displays attract mates. Or, he may have been threatening another individual if he was crouched over and pointing directly at another penguin. This is known as a Gape. It is still possible that this male or female was simply greeting his/her mate. Mates, after periods of absence, will approach each other and perform mutual displays and mutual trumpet displays. 20. How do penguins greet each other? penguins greet each other in a variety of ways. Sometimes this is dependent upon the species; other times it is not. After periods of prolonged absence, pairs usually greet each other by reinforcing the pair bond. This is usually accomplished by something called the "mutual display." The mutual display sounds like and ecstatic display, although it's not initiated with as much effort. Both penguins engage in the act and are frequently facing each other when it occurs. Other penguins may only grunt or do nothing at all. A common practice among the Spheniscids is "billing." Pairs approach each other and tap their bills against each other. 21. Do penguins ever fight? Certainly. Penguins fight as much as other animals, and sometimes these fights can be fairly intense. Fighting is usually most intense around the mating season. Depending on the location and species, fighting can vary from wing tapping and bickering to seemingly hateful jabbing, biting, kicking and chasing across the colony. Penguins like the King and Emperor don't lend themselves to aggression all that often. For Emperors especially, fighting during mating season is particularly expensive. Penguins like Fiordlands, Macaronis and Rockhoppers get fairly rowdy with each other. I spent a few winters observing Humboldt penguins. I found them to be fairly amicable species that fought only rarely. They engaged in what is called "binary exchanges." This means that there were only two acts to aggression. The first act was the first aggressive act and the second act was the response to the first act. Usually the second act, or response, was to retreat or turn away from the aggression. This indicates that fighting was either not economically feasible at the point or that the fight ended with submission to the dominant individual. However, frequency of fighting and intensity (more acts) of fighting occurred around mating season. 22. How do penguins reproduce? Penguins reproduce in a fairly simply way. Reproduction is by sexual means (i.e., there is the exchange of genetic information via sperm and ovum). There is fairly good development of this in the ethology and biology sections. 23. Are penguins intelligent? I'm afraid to say that penguins won't be solving algebra equations any time soon. They're probably not on the bright side of the classroom either. Experiments have shown that penguins are capable of basic Pavlovian training (e.g., reward given for following a simple command), but not much beyond that. They have been trained to stop on command and perform some of the simpler tasks domesticated dogs can do. That's about it. They're not very intelligent.

24. Do penguins mate for life? Most avian species are monogamous. That is to say that when they do mate, they only mate with one other. That doesn't imply that during the next mating season they will mate with the same individual. However, some birds do mate for life. Penguins are more-or-less monogamous, but some species are known to seek extra-pair copulations, that is, have affairs. The whole idea of mating for life is a bit exaggerated. 25. I heard penguins offer pebbles as engagement rings. Is this true? No. There has been at least one study of Adelie penguins in which the male offers pebbles (as part of the nest) to the female. It has been theorized that this "gift" is actually a demonstration to the female that the male is capable of building a nest and would -- by some property -- be a good mate and "father." This sort of "gift giving" isn't uncommon in animal and insect populations. This act certainly doesn't occur in all penguin species and isn't by any stretch an engagement gift. 26. How long do penguins live? Penguins in captivity will live longer than penguins in the wild since they are fed nutrionally balanced meals and do not have to contend with predators. Generally, penguins will live from 15 to 20 years. A female Humboldt penguin in my study group (in captivity) lived to 17 years. 27. Do penguins have knees? Yes. It doesn't look like they have much in the way of legs, but their knees are hidden under their feathers. Here's a basic diagram: Most people think that the knees on birds are "backwards." What you are actually seeing is a structure analagous to the human ankle.

28. Why don't penguin feet freeze after they leave the water? The system that enables a penguin foot to not freeze is basically the system that keeps your fingers from freezing while in the cold, only that system in the penguin is much more extensive and elaborated. Imagine a section of a penguin foot. Capillaries extend into ridges of penguin skin (see penguin histology). They feed oxygen directly to the remotest parts of the skin. At this point, blood is fed from the deeper plexuses and from the larger

arteries of the penguin vasculature. Ultimately, oxygen and heat supply to the skin is a function of cardiac output, or heart rate with respect to volume of blood pumped. The following diagram shows the network of blood vessels in that section.
A: Capillaries. B: Superficial plexus. C: Subsuperficial plexus. D: Intermediate plexus. E: Deep plexus

How does this system work? It's actually quite ingenius. The metabolic processes of the penguin -- that is, the chemical reactions that generate both energy and heat -- require oxygen. The vasculature of the penguin delivers oxygen from the lungs to the heart and eventually the rest of the body. This vasculature also delivers heat and keeps it within the body. If you look closely at the diagram, you can see loops at the top. These loops, along with loops in the deeper plexuses create what is called the counter-current exchange system. That is, when heat is lost from the arterial side of the capillaries, the venous side of the capillaries running in the opposite direction pick up that heat so it is not lost. This means that in the very remote regions of the skin, cells get oxygen but heat isn't lost through this skin. As a result, penguin feet don't freeze since heat is never lost. 29. Well, how do penguins cool off? This question may make sense for penguins that live in Australia and the Galapagos Islands, but for penguins in Antarctica? Yes, all penguins overheat from time to time. However, overheating is more of a problem for penguins living in semi-tropical regions like the Yellow-eyed penguin or the desert regions like that Humboldt and Magellanic penguin. The Spheniscid penguins, i.e. the Galapagos, Magellanic, Humboldt and African, have featherless patches around their heads. In these regions without feathers, air can't be trapped, and heat is given off rapidly at this point. Despite the fact that the counter current exchange is so efficient, some heat is lost in the counter current exchange when it is overloaded. Normally this heat is trapped in the feathers, but bare patches allow it to be given off. In areas where there are feathers, penguins contract muscles called feather erectors which cause their feathers to become erect. This eliminates a good deal of the covered insulation feathers supply and allows heat to exchange with the atmosphere. These mechanisms allow the penguin too cool down. Furthermore, they are under autonomic regulation -- that is, the penguin does this automatically without "thinking" about it. Additionally, the penguin air sacs form reservoirs of air that also permit the collection and dissipation of air through respiration. That's why you might see penguins with their bills wide open panting. This is partially conscious, but it is mostly autonomic as well. 30. Do any penguins live in the Arctic Circle? No. Penguins didn't evolve in the northern hemsiphere, and with the exception of the Galapagos penguin, no species of penguin naturally inhabits any land north of the equator. Galapagos penguins sometimes feed and stray slighly north of the equator.

31. I heard there was a mole that lived in the icy grounds of Antarctica that eats penguins. Is this true? No. It was published in an April Fool's edition of Discovery Magazine. No land mammals are indiginous to Antarctica. How silly!


Emperor penguin creches and altruism Kevin C Welch, 1999

What are penguin creches? That statement is a question which can be answered on several levels. The first level of analysis includes a descriptive and functional assay of penguin creches. Describing creches and demonstrating their purposes are subjects easily explored. The more interesting question - and perhaps the one with an almost unattainable answer -- is what are the evolutionary and behavioural purposes of a creche? The answer to that question defies current perception of animal altruism by the general public. Briefly, a creche is an aggregation or accumulation of penguins in close apposition or proximity, usually touching. Creches are found almost entirely in the emperor penguin populations, and they are formed by both adults and juveniles. The Antarctic climate is particularly hostile, especially in the depth of winter when temperatures can reach well below freezing. Coupled with brutal winds, the Antarctic shelves are areas in which few animals could survive. Emperor adults, while cradling their single eggs on their feet, form groups of tightly knit populations. Emperor penguin juveniles do the same thing once they are capable of being somewhat independent. The creche is composed of a periphery (i.e., the outside) and a central portion or inner range. Capillary loss of heat travels by convection in the creche to near-by penguins where it is often sequestered. Heat lost by penguins in the periphery is just that: lost. Penguins on the periphery lose more heat than they gain, while those nearest the center retain and receive more heat. Additionally, winds and drifting snow bombard members of the periphery accelerating the heat lost by these members. Wind minimally contacts central members of the creche. Throughout the nights and coldest days while these creches exist, members of the periphery move about to the central regions and the creche appears to move en masse from its original
formation spot -- gradually, that is. In summary, the creche serves as a means for staying warm. This addresses the first question.

The notion that individuals on the periphery take their turns on the outside and that members on the inside relinquish their turns at the center so that the two populations may "switch" is erroneous. It's a

fairly codified hypothesis of ethology and evolution that the rule of genetic fitness and "selfish genetics" (i.e., looking out for number one) is supreme. Individual penguins are simply not willing to take turns in any of these positions in the creche; each and every one is focused on getting to the center and remaining at the center simply because each is probably uninterested in the collective survival of the group. To suggest that emperor penguins take their turns so that other penguins have a chance of staying warm is otherwise known as group selection. Group selection is a largely discounted evolutionary theory that presupposes that individuals conduct themselves to ensure the survival of the group with the subtending notion that group survival leads to individual survival. A more appropriate and correct evolutionary theory is kin selection. Kin selection hypothesizes that individuals conduct themselves in such a manner that ensures a survival of that individual as well as those related to it. In the case of emperor juveniles, each chick is the sole offspring of its parents; therefore, the emperor juvenile has no kin, per se. Hence, the emperor juvenile looks out for itself, and it most likely makes every effort to attain a central position in the creche and keep it. This is why creches appear to move; each emperor juvenile is jockeying for an optimal position in the creche. This theory begets a few questions: if the center is such a prized location, why don't penguins fight for the center? If they are unwilling to fight, why? They're certainly valid questions and may even present caveats to the theory. Simply, fighting is always a last resort in animals. Most animals undergo extensive and ritualized displays to exert dominance before fighting. Most of these displays result in one of the two backing down. That solves part of the question. But, what about those willing to fight? With temperatures below -100 and the scarcity of food, fighting is expensive energetically and repair may not be possible under these winter conditions. With the risk of injury and loss of energy, the absence of fighting -- usually restricted to petty squabbling and flapping -- is observed since it is more appealing than a fight over a possibly better position that constantly changes. This does not, however, preclude attempts and foolery. Those that don't achieve central positions are more likely to succumb to the weather, a demonstration of natural selection.

Darwin, Charles. On Evolution. Glick & Kohn, eds. 1996. Hackett Publishing Company, Inc.: Indianapolis. Krebs, J.R. & Davies, N.B. 1993. An Introduction to Behavioural Ecology. London: Blackwell Scientific Publications. Krebs, J.R. & Davies, N.B. 1991. Behavioural Ecology: An Evolutionary Approach. London: Blackwell Scientific Publications. Young, David. 1992. The Discovery of Evolution. Cambridge Uniersity Press: Cambridge.