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Three years later, the great British ichthyologist C. Tate Regan, then keeper of fishes and later boss of the British Museum (Natural History), solved Saemundsson’s dilemma. The “young ones” were not juveniles, but permanently attached, sexually mature dwarf males. As Regan studied the details of attachment between male and female, he discovered the astounding fact that has ever since been celebrated as one of the greatest oddities in natural history: “At the junction of the male and the female fish there is a complete blending…their vascular systems are continuous.” In other words, the male has ceased to function as an independent organism. It no longer feeds, for its mouth is fused with the female’s outer skin. The vascular systems of male and female have united, and the tiny male is entirely dependent upon the female’s blood for nutrition. Of a second species with similar habits, Regan writes: “It is impossible to say where one fish begins and the other ends.” The male has become a sexual appendage of the female, a kind of incorporated penis. (Both popular and technical literature often refer to the fused male as a “parasite.” But I demur. Parasites live at the expense of their host. Fused males depend upon females for nutrition, but they supply in return that most precious of biological gifts—access to the next generation and a chance for evolutionary continuity.)

The extent of male submergence has been exaggerated in most popular accounts. Although attached males surrender their vascular independence and lose or reduce a set of organs no longer needed (eyes, for example), they remain more than a simple penis. Their own hearts must still pump the blood now supplied by females, and they continue to breathe with their gills and remove wastes with their kidneys. Of one firmly attached male, Regan writes:

The male fish, although to a great extent merely an appendage of the female, and entirely dependent on her for nutrition, yet retains a certain autonomy. He is probably capable, by movements of the tail and fins, of changing his position to some extent. He breathes, he may have functional kidneys, and he removes from the blood certain products of his own metabolism and keeps them as pigment…. But so perfect and complete is the union of husband and wife that one may almost be sure that their genital glands ripen simultaneously, and it is perhaps not too fanciful to think that the female may possibly be able to control the seminal discharge of the male and to ensure that it takes place at the right time for the fertilization of her eggs.

Nonetheless, however autonomous, the males have not honed themselves to Darwinian optimality, for they have evolved no mechanism for excluding other males from subsequent attachment. Several males are often embedded into a single female.

(While criticizing the exaggeration of some popular accounts, allow me a tangential excursion to express a pet peeve. I relied upon primary, technical literature for all my descriptions, but I began by reading several popular renditions. All versions written for nonscientists speak of fused males as the curious tale of
the
anglerfish—just as we so often hear about
the
monkey swinging through trees, or
the
worm burrowing through soil. But if nature teaches any lesson, it loudly proclaims life’s diversity. There ain’t no such abstraction as
the
clam,
the
fly, or
the
anglerfish. Ceratioid anglerfishes come in nearly 100 species, and each has its own peculiarity. Fused males have not evolved in all species. In some, males attach temporarily, presumably at times of spawning, but never fuse. In others, some males fuse and others become sexually mature while retaining their bodily independence. In still others, fusion is obligatory. In one species of obligate fusers, no sexually mature female has ever been found without an attached male—and the stimulus provided by male hormones may be a prerequisite for maturation.

These obligate fusers have become the paradigm for popular descriptions of
the
anglerfish, but they do not represent the majority of ceratioid species. I grouse because these meaningless abstractions convey seriously false impressions about nature. They greatly exaggerate nature’s discontinuities by focusing on extreme forms as false paradigms for an entire group, and rarely mentioning the structurally intermediate species that often live happily and abundantly. If all fishes either had totally independent or completely fused males, then how could we even imagine an evolutionary transition to the peculiar sexual system of
the
anglerfish? But the abundance of structurally intermediate stages—temporary attachment or fusion of some males only—conveys an evolutionary message. These modern structural intermediates are not, of course, actual ancestors of fully fused species, but they do sketch an evolutionary pathway—just as Darwin studied the simple eyes of worms and scallops to learn how a structure so complex and apparently perfect as the vertebrate eye might evolve through a chain of intermediate forms. In any case, bursting diversity is nature’s watchword; it should never be submerged by careless abstraction.)

Ceratioid males embark upon their peculiar course early in life. As larvae, they feed normally and live independently. After a period of rapid change, or metamorphosis, males in species destined for fusion do not develop their alimentary canals any further, and never feed again. Their ordinary teeth disappear, and they retain and exaggerate only a few fused teeth at the tips of their mouth—useless in feeding, but well adapted for piercing and holding tight to a female. They become sleek and more streamlined, with a pointed head, compressed body, and strong, propulsive tail fin—in short, a sort of sexual torpedo.

But how do they find females, those tiny dots of connubial matter in the midst of an endless ocean? Most species must use olfactory cues, a system often exquisitely developed in fishes, as in homing salmon that smell out their natal stream. These ceratioid males develop gigantic nostrils after metamorphosis; relative to body size, some ceratioids have larger nasal organs than any other vertebrate. Another family of ceratioids fails to develop large nostrils, but these males have enormously enlarged eyes, and they must search for the ghostly light of fishing females (each species has a different pattern of illumination, and males probably recognize their proper females). The system is not entirely failsafe, as ichthyologist Ted Pietsch recently found a male of one species attached to a female of a different species—a fatal mistake in evolutionary terms (although the two fish had not fused and might later have separated had not zealous science found and preserved them
in flagrante delicto
).

As I sit here wiggling my toes and flexing my fingers in glorious independence (and with a full one-inch advantage over my wife), I am tempted (but must resist) to apply the standards of my own cherished independence and to pity the poor fused male. It may not be much of a life in our terms, but it keeps several species of anglerfishes going in a strange and difficult environment. And who can judge anyway? In some ultimate Freudian sense, what male could resist the fantasy of life as a penis with a heart, deeply and permanently embedded within a caring and providing female? These anglerfishes represent, in any case, only the extreme expression of nature’s more common pattern—smaller males pursuing an evolutionary role as sources of sperm. Do they not, therefore, teach us a generality by their very exaggeration of it? We human males are the oddballs.

I therefore take my leave of fused anglerfishes with a certain sense of awe. Have they not discovered and irrevocably established for themselves what, according to Shakespeare, “every wise man’s son doth know”—“journeys end in lovers meeting”?

2 | Nonmoral Nature

WHEN THE
Right Honorable and Reverend Francis Henry, earl of Bridgewater, died in February, 1829, he left £8,000 to support a series of books “on the power, wisdom and goodness of God, as manifested in the creation.” William Buckland, England’s first official academic geologist and later dean of Westminster, was invited to compose one of the nine Bridgewater Treatises. In it he discussed the most pressing problem of natural theology: if God is benevolent and the Creation displays his “power, wisdom and goodness,” then why are we surrounded with pain, suffering, and apparently senseless cruelty in the animal world?

Buckland considered the depredation of “carnivorous races” as the primary challenge to an idealized world where the lion might dwell with the lamb. He resolved the issue to his satisfaction by arguing that carnivores actually increase “the aggregate of animal enjoyment” and “diminish that of pain.” Death, after all, is swift and relatively painless, victims are spared the ravages of decrepitude and senility, and populations do not outrun their food supply to the greater sorrow of all. God knew what he was doing when he made lions. Buckland concluded in hardly concealed rapture:

The appointment of death by the agency of carnivora, as the ordinary termination of animal existence, appears therefore in its main results to be a dispensation of benevolence; it deducts much from the aggregate amount of the pain of universal death; it abridges, and almost annihilates, throughout the brute creation, the misery of disease, and accidental injuries, and lingering decay; and imposes such salutary restraint upon excessive increase of numbers, that the supply of food maintains perpetually a due ratio to the demand. The result is, that the surface of the land and depths of the waters are ever crowded with myriads of animated beings, the pleasures of whose life are coextensive with its duration; and which throughout the little day of existence that is allotted to them, fulfill with joy the functions for which they were created.

We may find a certain amusing charm in Buckland’s vision today, but such arguments did begin to address “the problem of evil” for many of Buckland’s contemporaries—how could a benevolent God create such a world of carnage and bloodshed? Yet this argument could not abolish the problem of evil entirely, for nature includes many phenomena far more horrible in our eyes than simple predation. I suspect that nothing evokes greater disgust in most of us than slow destruction of a host by an internal parasite—gradual ingestion, bit by bit, from the inside. In no other way can I explain why
Alien
, an uninspired, grade-C, formula horror film, should have won such a following. That single scene of Mr. Alien, popping forth as a baby parasite from the body of a human host, was both sickening and stunning. Our nineteenth-century forebears maintained similar feelings. The greatest challenge to their concept of a benevolent deity was not simple predation—but slow death by parasitic ingestion. The classic case, treated at length by all great naturalists, invoked the so-called ichneumon fly. Buckland had sidestepped the major issue.

The “ichneumon fly,” which provoked such concern among natural theologians, was actually a composite creature representing the habits of an enormous tribe. The Ichneumonoidea are a group of wasps, not flies, that include more species than all the vertebrates combined (wasps, with ants and bees, constitute the order Hymenoptera; flies, with their two wings—wasps have four—form the order Diptera). In addition, many non-ichneumonid wasps of similar habits were often cited for the same grisly details. Thus, the famous story did not merely implicate a single aberrant species (perhaps a perverse leakage from Satan’s realm), but hundreds of thousands—a large chunk of what could only be God’s creation.

The ichneumons, like most wasps, generally live freely as adults but pass their larval life as parasites feeding on the bodies of other animals, almost invariably members of their own phylum, the Arthropoda. The most common victims are caterpillars (butterfly and moth larvae), but some ichneumons prefer aphids and others attack spiders. Most hosts are parasitized as larvae, but some adults are attacked, and many tiny ichneumons inject their brood directly into the egg of their host.

The free-flying females locate an appropriate host and then convert it to a food factory for their own young. Parasitologists speak of ectoparasitism when the uninvited guest lives on the surface of its host, and endoparasitism when the parasite dwells within. Among endoparasitic ichneumons, adult females pierce the host with their ovipositor and deposit eggs within. (The ovipositor, a thin tube extending backward from the wasp’s rear end, may be many times as long as the body itself.) Usually, the host is not otherwise inconvenienced for the moment, at least until the eggs hatch and the ichneumon larvae begin their grim work of interior excavation.

Among ectoparasites, however, many females lay their eggs directly upon the host’s body. Since an active host would easily dislodge the egg, the ichneumon mother often simultaneously injects a toxin that paralyzes the caterpillar or other victim. The paralysis may be permanent, and the caterpillar lies, alive but immobile, with the agent of its future destruction secure on its belly. The egg hatches, the helpless caterpillar twitches, the wasp larva pierces and begins its grisly feast.

Since a dead and decaying caterpillar will do the wasp larva no good, it eats in a pattern that cannot help but recall, in our inappropriate, anthropocentric interpretation, the ancient English penalty for treason—drawing and quartering, with its explicit object of extracting as much torment as possible by keeping the victim alive and sentient. As the king’s executioner drew out and burned his client’s entrails, so does the ichneumon larva eat fat bodies and digestive organs first, keeping the caterpillar alive by preserving intact the essential heart and central nervous system. Finally, the larva completes its work and kills its victim, leaving behind the caterpillar’s empty shell. Is it any wonder that ichneumons, not snakes or lions, stood as the paramount challenge to God’s benevolence during the heyday of natural theology?

As I read through the nineteenth- and twentieth-century literature on ichneumons, nothing amused me more than the tension between an intellectual knowledge that wasps should not be described in human terms and a literary or emotional inability to avoid the familiar categories of epic and narrative, pain and destruction, victim and vanquisher. We seem to be caught in the mythic structures of our own cultural sagas, quite unable, even in our basic descriptions, to use any other language than the metaphors of battle and conquest. We cannot render this corner of natural history as anything but story, combining the themes of grim horror and fascination and usually ending not so much with pity for the caterpillar as with admiration for the efficiency of the ichneumon.

I detect two basic themes in most epic descriptions: the struggles of prey and the ruthless efficiency of parasites. Although we acknowledge that we may be witnessing little more than automatic instinct or physiological reaction, still we describe the defenses of hosts as though they represented conscious struggles. Thus, aphids kick and caterpillars may wriggle violently as wasps attempt to insert their ovipositors. The pupa of the tortoiseshell butterfly (usually considered an inert creature silently awaiting its conversion from duckling to swan) may contort its abdominal region so sharply that attacking wasps are thrown into the air. The caterpillars of
Hapalia
, when attacked by the wasp
Apanteles machaeralis
, drop suddenly from their leaves and suspend themselves in air by a silken thread. But the wasp may run down the thread and insert its eggs nonetheless. Some hosts can encapsulate the injected egg with blood cells that aggregate and harden, thus suffocating the parasite.

J. H. Fabre, the great nineteenth-century French entomologist, who remains to this day the preeminently literate natural historian of insects, made a special study of parasitic wasps and wrote with an unabashed anthropocentrism about the struggles of paralyzed victims (see his books
Insect Life
and
The Wonders of Instinct
). He describes some imperfectly paralyzed caterpillars that struggle so violently every time a parasite approaches that the wasp larvae must feed with unusual caution. They attach themselves to a silken strand from the roof of their burrow and descend upon a safe and exposed part of the caterpillar:

The grub is at dinner: head downwards, it is digging into the limp belly of one of the caterpillars…. At the least sign of danger in the heap of caterpillars, the larva retreats…and climbs back to the ceiling, where the swarming rabble cannot reach it. When peace is restored, it slides down [its silken cord] and returns to table, with its head over the viands and its rear upturned and ready to withdraw in case of need.

In another chapter, he describes the fate of a paralyzed cricket:

One may see the cricket, bitten to the quick, vainly move its antennae and abdominal styles, open and close its empty jaws, and even move a foot, but the larva is safe and searches its vitals with impunity. What an awful nightmare for the paralyzed cricket!

Fabre even learned to feed paralyzed victims by placing a syrup of sugar and water on their mouthparts—thus showing that they remained alive, sentient, and (by implication) grateful for any palliation of their inevitable fate. If Jesus, immobile and thirsting on the cross, received only vinegar from his tormentors, Fabre at least could make an ending bittersweet.

The second theme, ruthless efficiency of the parasites, leads to the opposite conclusion—grudging admiration for the victors. We learn of their skill in capturing dangerous hosts often many times larger than themselves. Caterpillars may be easy game, but psammocharid wasps prefer spiders. They must insert their ovipositors in a safe and precise spot. Some leave a paralyzed spider in its own burrow.
Planiceps hirsutus
, for example, parasitizes a California trapdoor spider. It searches for spider tubes on sand dunes, then digs into nearby sand to disturb the spider’s home and drive it out. When the spider emerges, the wasp attacks, paralyzes its victim, drags it back into its own tube, shuts and fastens the trapdoor, and deposits a single egg upon the spider’s abdomen. Other psammocharids will drag a heavy spider back to a previously prepared cluster of clay or mud cells. Some amputate a spider’s legs to make the passage easier. Others fly back over water, skimming a buoyant spider along the surface.

Some wasps must battle with other parasites over a host’s body.
Rhyssella curvipes
can detect the larvae of wood wasps deep within alder wood and drill down to a potential victim with its sharply ridged ovipositor.
Pseudorhyssa alpestris
, a related parasite, cannot drill directly into wood since its slender ovipositor bears only rudimentary cutting ridges. It locates the holes made by
Rhyssella
, inserts its ovipositor, and lays an egg on the host (already conveniently paralyzed by
Rhyssella
), right next to the egg deposited by its relative. The two eggs hatch at about the same time, but the larva of
Pseudorhyssa
has a bigger head bearing much larger mandibles.
Pseudorhyssa
seizes the smaller
Rhyssella
larva, destroys it, and proceeds to feast upon a banquet already well prepared.

Other praises for the efficiency of mothers invoke the themes of early, quick, and often. Many ichneumons don’t even wait for their hosts to develop into larvae, but parasitize the egg directly (larval wasps may then either drain the egg itself or enter the developing host larva). Others simply move fast.
Apanteles militaris
can deposit up to seventy-two eggs in a single second. Still others are doggedly persistent.
Aphidius gomezi
females produce up to 1,500 eggs and can parasitize as many as 600 aphids in a single working day. In a bizarre twist upon “often,” some wasps indulge in polyembryony, a kind of iterated supertwining. A single egg divides into cells that aggregate into as many as 500 individuals. Since some polyembryonic wasps parasitize caterpillars much larger than themselves and may lay up to six eggs in each, as many as 3,000 larvae may develop within, and feed upon a single host. These wasps are endoparasites and do not paralyze their victims. The caterpillars writhe back and forth, not (one suspects) from pain, but merely in response to the commotion induced by thousands of wasp larvae feeding within.

Maternal efficiency is often matched by larval aptitude. I have already mentioned the pattern of eating less essential parts first, thus keeping the host alive and fresh to its final and merciful dispatch. After the larva digests every edible morsel of its victim (if only to prevent later fouling of its abode by decaying tissue), it may still use the outer shell of its host. One aphid parasite cuts a hole in the bottom of its victim’s shell, glues the skeleton to a leaf by sticky secretions from its salivary gland, and then spins a cocoon to pupate within the aphid’s shell.

In using inappropriate anthropocentric language for this romp through the natural history of ichneumons, I have tried to emphasize just why these wasps became a preeminent challenge to natural theology—the antiquated doctrine that attempted to infer God’s essence from the products of his creation. I have used twentieth-century examples for the most part, but all themes were known and stressed by the great nineteenth-century natural theologians. How then did they square the habits of these wasps with the goodness of God? How did they extract themselves from this dilemma of their own making?

The strategies were as varied as the practitioners; they shared only the theme of special pleading for an a priori doctrine—our naturalists
knew
that God’s benevolence was lurking somewhere behind all these tales of apparent horror. Charles Lyell, for example, in the first edition of his epochal
Principles of Geology
(1830–1833), decided that caterpillars posed such a threat to vegetation that any natural checks upon them could only reflect well upon a creating deity, for caterpillars would destroy human agriculture “did not Providence put causes in operation to keep them in due bounds.”

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