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Developmental stages of a haemogregarine were found within polychromatocytes and erythrocytes in Giemsa-stained blood smears from six evileye pufferfish (Amblyrhynchotes honckenii) caught at Koppie Alleen in the De Hoop Nature Reserve, South Africa. This unusual haemogregarine, Haemogregarina (sensu lato) koppiensis sp. nov., was characterized by encapsulated gamonts with recurved tails, features more common in haemogregarines infecting amphibian and reptilian erythrocytes than in those from fish. Haemogregarina koppiensis is only the third species of fish haemogregarine to have been described from South Africa.
Extensive field observations were made in the Mingan Islands, northern Gulf of St Lawrence, to evaluate spatial and temporal variations in the use of habitat and prey resources by two major subtidal predators, the seastars Leptasterias polaris and Asterias vulgaris. Although both seastars have similar size structures and generally overlapped in their spatial and temporal distribution, the degree of overlap varied in different sites and appeared to be related to prey abundance, substratum type and slope. Three general patterns were observed: (1) both species aggregating in shallow water and decreasing in numbers with depth; (2) the two species showing inverse depth distributions; and (3) both seastars occurring in low numbers across the subtidal zone. Temporal changes in availability of the preferred prey of the two seastars, the mussel Mytilus edulis, appeared to be a major factor affecting their abundance and distribution. The two seastars occurred together in dense aggregations on mussel beds in shallow water. They consume similar-sized mussels until the number of mussels becomes reduced when A. vulgaris, but not L. polaris, begins to select larger mussels. Once a mussel bed is decimated, the seastars appear to move away, possibly in search of other beds. Intensive seastar foraging limits the distribution of mussels to a few metres in depth. Below the mussel zone, the two seastars are spatially segregated at a small spatial scale (1 m2 quadrat) and select different alternative prey, L. polaris feeding mainly on the crevice-dwelling clam Hiatella arctica and A. vulgaris on the ophiuroid Ophiopholis aculeata. The size partitioning of the preferred prey in shallow water, and spatial segregation and selection of different alternative prey at greater depths may reflect mechanisms permitting the two seastars to coexist.
A feeding experiment was conducted on the marine worm Nereis diversicolor to compare the fate of a hydrocarbon mixture during the gut passage in this species with the hydrocarbon breakdown process demonstrated for Nereis virens. Hydrocarbon dissolution/solubilization processes in the gut of N. diversicolor were found to have similar qualitative and quantitative importance in the hydrocarbon transformation as those observed in N. virens.
Burrow morphology and mating behaviour of Upogebia noronhensis was studied using resin casting of burrows in the field and observation of laboratory maintained animals. Burrows of U. noronhensis showed a typical Y-shaped pattern in over 70% of the cases analysed. The remaining 30% comprised U-shaped burrows lacking the lower tunnel (shaft) and burrows with long additional branches projecting from the U portion. Results from animals left to construct burrows in the aquarium closely matched those found in nature. Field and laboratory burrows showed that different shapes are related to the collapse of the burrow walls, the burrowing activities of other individuals and species, and to the behaviour of the species itself. U-shaped burrows form as a result of the partial construction of the burrow (the U part is always built first) or owing to the collapse of the shaft. Burrows with additional branches always belonged to males and result from their search for a female with which to reproduce. This process also produced connected burrows. Mating occurs within the female burrow and this is the only time when two animals occupy the same burrow. After mating, the male returns to its burrow and immediately closes the connection. Larvae are planktonic and probably settle in adult areas, since the smallest juvenile burrows were always associated with adult burrows. This may contribute to the high population densities found in the field (∼200 ind m−2), which in consequence facilitates fossorial encounters for reproduction. This is the first time fossorial encounters for reproduction are reported for an Upogebia species and probably for all Thalassinidea.
The annual pattern of burial depth in natural populations of the infaunal bivalves Tagelus dombeii (Tellinacea) and Venus antiqua (Veneracea) is described in relation to annual food availability in both the water column and the sediment and abiotic factors (temperature and salinity) at Coihuín tidal flat, in southern Chile. A field experiment in which burial depth was measured in situ each month (over 14 months), with the aid of a fixed-length nylon thread attached to the shell. For T. dombeii the results showed a significant increase in burial depth with increasing bivalve size and syphon weight. Tagelus dombeii had a mean burial depth of 17·5 cm, which was three times more than in V. antiqua (5·30 cm). The burial depth dynamics for both species displayed a strong correlation with food availability in the water column. Approximately 60% of the variability in burial depth in T. dombeii and V. antiqua was explained by concentration of chlorophyll-a in the water column. Food concentration on the sediment surface did not effect burial depth, i.e. deposit feeding seems to be of minor significance in either species.
Large numbers of small individuals (pediveligers and juveniles <5 mm) are routinely recorded in size–frequency distributions of mussel samples collected from deep-sea chemosynthetic environments. If recruitment of invertebrates to deep-sea hydrothermal vent sites were via long-distance dispersal, as is typically assumed, one would expect recruitment ‘events’ recorded in size–frequency distributions to be difficult to detect, due to loss of larvae in an open system over large distances. If one imposes mesoscale oceanographic phenomena that minimize dilution of larvae (such as eddies shed from hydrothermal vent plumes) and episodic spawning, expression of this mesoscale corralling at the level of population structure would likely be limited to discrete records of recruitment events encountered serendipitously during haphazard sampling in space and time. The ubiquity of large numbers of post-larvae in mussel samples from a number of disparate sites is likely not serendipitous, but instead may reflect the importance of local sources and sinks of propagules in maintenance of mussel populations.
Specimens of the chemoautotrophic symbiont-bearing hydrothermal vent clam Calyptogena pacifica were collected from hydrothermal vents at the Endeavour segment of the Juan de Fuca Ridge. Total lipid was extracted from gill, foot and mantle tissues, and lipid class and fatty acid composition determined by thin layer chromatography with flame ionization detection (TLC–FID), gas chromatography (GC) and gas chromatography with mass spectrometry (GC–MS). An abundance of n–7 monounsaturated fatty acids (MUFA), especially in the gill, reflected the large contribution of chemoautotrophic symbiotic bacteria to the nutrition of this clam. The absence of n–8 MUFA suggests that C. pacifica does not contain methanotrophic symbiotic bacteria. Low levels of highly unsaturated fatty acids (HUFA) such as 20:5 n–3 and 22:6 n–3 were detected in C. pacifica and their presence is attributed to a source other than chemoautotrophic symbiotic bacteria. Significant levels of non-methylene interrupted dienoic fatty acids and eicosatrienoic acid (20:3) were also detected in C. pacifica and it is suggested that these fatty acids are synthesized from n–7 MUFA as alternatives to HUFA. In contrast to shallow water bivalves, elevated levels of triglyceride were detected in the gills compared to the mantle.
Virus-like particles (VLPs) were observed in thin sections of the plumose anemone, Metridium senile, collected from seawater off the west coast of Sweden in 1969. The VLPs were observed in the nucleus of spiroblasts and amoebocytes; they were either pentagonal or hexagonal in section, indicating an icosahedral structure. Virus-like particles were ∼60 nm in diameter, with an electron-dense core 40 nm in width; they had no apparent tail. This is the first substantial report of viruses observed in sea anemone cells.
Length–weight relationships were estimated for 18 Red Sea immigrant fish species from the eastern Mediterranean coasts of Turkey. The values of the exponent b in the length–weight regression (W= aLb) ranged between 2·482 and 3·355. The median value was 3·018 and 50% of the values ranged between 2·835 and 3·121.
Preferences of the sea urchin Diadema antillarum (Echinodermata: Echinoidea) feeding on five species of brown macroalgae (Padina pavonica, Dyctiota dychotoma, Cystoseira abies-marina, Lobophora variegata and Halopteris filicina) have been studied using caging field experiments on Gran Canaria Island during August to October 2000. Results of three assays of both single and multiple diet experiments rejected the null hypothesis that Diadema does not feed selectively on the five selected algal species. In the multiple diet assays, Diadema consumed an average of 68–98 mg algae urchin−1 h−1 and 4–120 mg algae urchin−1 h−1 in the single diet experiments. On the basis of consumption, the five species of algae eaten can be divided into three groups. Thus Halopteris, Lobophora and Dyctiota were considered preferred algae, while Padina was considered an intermediate alga. Finally Cystoseira was significantly the less preferred and consumed seaweed in all sets of assays.
Modern humans stayed anatomically unchanged at least for the past 80,000 years. On the evolutionary level of organization, anatomically fixed things are expected to stay (nearly) fixed in behavior. Our (anatomically) modern ancestors lived up to this rule for the first half, or slightly more, of their tenure on earth. All hell broke loose in the second. The extraordinary changes in the archeological record starting around 40,000 to 30,000 years ago, and carrying through the height of the last ice age to the onset of the Holocene (some 10,000 years ago), suggest remarkable refinements in behavioral structures unexpected of a morphologically fixed organism. Changes in the record further suggest a remarkable increase in regional and temporal diversity of material structures that up to that point varied little through time and space. The Middle to Upper Paleolithic transition, or the creative explosion as this episode has sometimes been labeled (e.g., Pfeiffer, 1982), is most vividly evident in wall paintings preserved in caves, in portable art, personal ornamentation, and in elaborate burials. More subtle are the sudden refinements in tools, and the rapid expansion into new geographic areas, indeed, into two new continents (Australia and the Americas). Underlying all of this is an authentic economic expansion reminiscent of various mercantile and industrial revolutions in recorded history.
The key question, from an evolutionary viewpoint, is how could such remarkable changes take place in functional behavior without apparent change in morphology. One possible explanation ascribes this turn of events to some neurological change that led to an evolution in behavior without an apparent change in anatomical form (e.g., Klein 1992).
Among the sailors who crisscrossed the oceans during the sixteenth century few knew how to swim. It was pointless. A sailboat at sea could not be stopped or turned to rescue a shipmate washed or fallen overboard. The hazards of the occupation could not escape the attention of sailors struggling to lash unruly sails in unpredictable gale-force winds. One slip on the upper yards meant almost certain death. Given the state of the art and the perils of the sea, casualties from accidents and disease were exceedingly high even by the standards of the time. As to comforts and niceties of life – nobody could ask for, nor possibly be granted, accommodations beyond the bare minimum.
With lives aboard more expendable than cargo, one can wonder why skippers and ship stewards paid meticulous attention to supplies of food-stuff. A typical list of ship supplies for a journey across the Atlantic around 1577, the year Sir Francis Drake set out to circumnavigate the world, included such diverse items as hardtack, flour, pickled beef and pork, dried peas and codfish, butter, cheese, oatmeal, rice, honey, and vinegar, as well as about eight tons of beer. The selection seems to compare quite favorably even today with any decent hospital or school cafeteria – not counting the beer. Careful attention to an adequate bulk of food supplies before embarking on unpredictably long trips such as the one undertaken by the Mayflower (66 days across the Atlantic in 1620) is perfectly understandable. But why overburden the already hard-pressed logistics of a ship on high seas with a wide variety of food items?
Darwin and the Scottish economists: The first point of junction
The fundamental economic problem of human evolution
In view of the discussion thus far, the issue is no longer the mere existence of a human predisposition to exchange, but its evolutionary origin. Unsure of this origin, Adam Smith himself acknowledged (and deflected) in passing an intriguing question:
Whether this propensity [i.e., to exchange] be one of those original principles in human nature, of which no further account can be given; or whether, as seems more probable, it be the necessary consequence of the faculties of reason and speech, it belongs not to our present subject to enquire. (1976:17, italics added)
The inception of modern economics was thus accompanied (in 1776) by an evolutionary question preceding by nearly a century the Darwinian notion of evolution itself. Adam Smith probably agonized over this question, though as it seems, had the good sense to abort it in due course. Neither he nor his pre-Darwinian readers could fully comprehend the question, let alone conceive a sensible answer to it. With the advantage of hindsight, however, the question seems to present a challenge of the highest order to the modern study of human evolution: Was exchange an early agent of human evolution, or was it merely a late by-product of previously evolved “faculties of reason and speech”?
With the publication of The Descent of Man a century or so later, Darwin had at his disposal a fairly mature notion of evolution applicable to human affairs.
The first appearance of manufactured stone tools in widespread systematic human use, about 2 million years ago, in combination with other evidence (e.g., food sharing), suggests the possibility that some form of exchange, however primitive, was already practiced by traders with brains half the size of a modern human being. If true, the implication is that exchange could have played an important facilitating role already in the first major economic transition in human (or protohuman) history – the transition from the feed-as-you-go strategy of primate feeding to hunting-gathering. The discussion in this chapter will consider these possibilities in light of the available paleoarcheological data, starting with a preliminary review of the physical environment.
The physical environment
The single most important factor that describes the theater of events in evolution, certainly in human evolution, is climate. The global climate in which civilization flourished is the wrong environmental model for understanding human evolution not only throughout the long ice ages (nearly 90% of the time humans walk the earth) but, in all likelihood, also in the course of the short spikes of interglacial periods (the remaining 10%). The climate observed today and enjoyed by (anatomically) modern people over the past 10,000 years is a placidly warmand, arguably, all too brief interval in a long process of falling temperatures starting about 3.2 million years ago (and greatly intensifying roughly 2.4 million years ago and then again about 900,000 years ago).
If market exchange evolved from some preadaptation observable in humans or animals, then a careful examination of analogies with nepotistic exchange (such as courtship feeding in birds) or symbiotic exchange (such as pollination) is a logical course of action to pursue. In following this approach the first part of this chapter demonstrates the existence of some tempting analogies of market exchange drawn from the behavior of certain animals and plants, as well as from the nonmarket sphere of human affairs. It also demonstrates the risk of drawing premature, if not fanciful, explanations from such analogies.
Alternatively, if market exchange evolved de novo, then the major thrust in the inquiry is best directed toward the mechanisms of the market and the deep structures of exchange itself. This approach is undertaken in the second part of this chapter with the hope that intricacies of market exchange as we know them could provide a clue to their origin. At issue is a catalyst in the form of an activity, or perhaps a single commodity, with the power of spurring exchange between exceedingly reluctant traders at some remote point in antiquity.
Bateman's syndrome
There are certain parallels between the choice of mating partners and the choice of trading partners – chief among them is asymmetry. The asymmetry in the case of sexual selection is due to adaptive pressures that for a widely recognized reason – known as Bateman's principle – act differently on the reproductive behavior of males and females. In a pattern that cuts across nearly all the species of higher organisms, as already noted by Darwin (1874), females are more selective.
Without really being aware of it, human beings economically interact under two distinct routines – nepotistic exchange and market exchange – effortlessly switching back and forth between the two regimes. The tendency to turn on and off two separate processes of decision-making (with implications in action and in demeanor) could have been readily diagnosed as some sort of split-personality disorder had it not been so predictable – repeating itself in nearly the same manner in almost any member of the species. The fundamental difference between the market and the nepotistic (or domestic) routines of human activities was examined in the foregoing discussion (especially in Chapter 2) on the grounds of practical considerations and, so far, only with a secondary emphasis on evolutionary ones. The main implication suggests nothing in the way of a behavioral disorder. What it suggests is the coexistence of two distinct adaptations. The main attempts to trace these two adaptations to their separate evolutionary origins are relegated to two chapters in this volume. The present chapter is primarily concerned with the origins of nepotistic exchange. (Chapter 9 will be similarly concerned with the origins of market exchange.)
Primordial exchange at the lowest levels of organization
Certain advantages of division of labor, supported by exchange, occurred at fairly early stages of life on earth. Consider, for instance, the evolution of one of the most comprehensive and amazing systems of exchange and division of labor in nature: a single body of a multicelled organism. Finely tuned exchange among specialized cells, tissues, and organs is inherent in the physiology of such an organism.
The propensity and capacity to exchange one thing for another between two traders – however unrelated to each other – is a profound distinguishing feature of human subsistence. Human beings are endowed with remarkable skills of trade which they deploy spontaneously when confronted with favorable opportunities; skills that lie dormant in the absence of such opportunities. As is true of other innate human abilities – such as the mastery of spoken language – basic skills of trade are taken for granted precisely because they are either inborn or acquired at a young age without conscious effort. Such skills are not as trivial as they may seem to a casual observer or, for that matter, to their very practitioners. Exchange requires certain levels of dexterity in communication, quantification, abstraction, and orientation in time and space – all of which depend (i.e., put selection pressure) on the lingual, mathematical, and even artistic faculties of the human mind. Moreover, exchange relies on mutual trust: predictable codes of conduct agreeable to the human sense of morality. Exchange, therefore, is a pervasive human predisposition with obvious evolutionary implications. The root cause of this predisposition and its evolutionary consequences in history, and prehistory, are the central concerns of this book.
Was exchange an early agent of human evolution, or is it a mere de novo artifact of modern civilization? The evolutionary literature treats the question with great caution. Many authors, starting with Charles Darwin and Alfred Russel Wallace, preferred to avoid the issue altogether. When the issue comes to the fore, the importance of exchange in recent industrialized societies is readily acknowledged.