Bijdragen tot de Dierkunde, 60 (3/4) 151-154 (1990)

SPB Academie Publishing bv, The Hague

Frustrating facts about area cladistics and individuality

E. Gittenberger

P.O. Box NL 2300 RA Leiden, The Netherlands Rijksmuseum van Natuurlijke Historie, 9517,

Aciculidae Keywords: cladistics, area cladogram, ,

Abstract matism. Sometimes not the individualistic biologi-

in cal properties of a (group of) species existing na-

decisive for the who Area cladistics suffer from the fact that areas have various “his- ture seem to be biogeographer

such dictated different of taxa. Conse- tories”, as by groups has to explain a certain pattern, but his favorite ex-

quently,a generalizedarea cladogramhas not or hardlyany bio- planatory model. There are vicariantists, disper- logical meaning. Conflicting area cladograms may be derived salists and several other kinds of biogeographers in different from even the same species, when their ranges geo- biased look influenced with a at nature, primarily logicalperiods are used. This is a consequence of the individuali- the of their school instead of the facts ty of species. by dogmas by

of nature. As I tried to show before (Gittenberger,

1984), our objects of research, the biological spe- Résumé

cies, should be looked upon as either vicariantists,

science L’approche connuesous le nom d’“Area cladistics” souffre du dispersalists, or something in between. In

fait les aréaux semblent avoir des histoires différentes, dic- que there should be simply biogeographers. In a way

tées l’existence de différents de Par consé- par groupes taxa. object and subject have to be interchanged again.

quent, les “generalized area cladograms” n’ont pas (ou ont à

Des “area peine) une signification biologique. cladograms”

contradictoires peuvent être construits même en utilisant les

mêmes espèces, si leurs distribution dans des périodes géolo-

Tout giques différentes est prise en considération. ceci est une Good and bad conséquence de l’individualité des espèces. species

Introduction The present situation results in doomed species,

those neglected by the adherents of a certain school,

The present wealth in biogeographic theoretics and because they fit better into the modelof the compet-

modelling (see e.g. Myers & Giller (1988) and vari- ing biogeographers. Another consequence is the

be weird distributional ous articles in Systematic Zoology 37 (3-4)) can tendency to neglect patterns envisaged as token of a flourishing science, as well that do not fit nicely into any model. Apparently, as illustrative of Edington's rule that the number of there should be law and order in nature, good spe- different hypotheses erected to explain a given bio- cies and bad species, those illustrating general pat-

is aberrant. The latter logical phenomenon inversely proportional to the terns and those only category available knowledge. The actual development is condemned to obscurity by those who consider seems to be parallelized by an increasing dog- narratives inferior to scientific law. 152 E. Gittenberger - Area cladistics and species individuality

point of view of biologists equalizing history with

biological history, areas have nothing comparable.

Recently Cracraft (1988: 221) made a start in under-

mining the foundations of area cladistics, em-

phasizing that "areas can have multiple histories,

but taxa cannot". There is only one single phylo-

of the that genetic tree for a group taxa, one repre-

sents their phylogenetic history correctly. There are

for of of several area cladograms a group fragments

earth crust, depending upon the taxa with common

histories used to derive them. It can be of biological

and geological interest to trace these cladograms,

because they tell parts of a real story. However,

sim- constructing a generalized area cladogram by

ply putting everything together and calculating the

mean, is only a mathematical exercise.

General and special biogeography

Fig. 1. Miocene (M), Pleistocene (P), Holocene (H) and recent In fact, general biogeography, applying to the dis- (dots) records ofPlatylapolita (Hartmann) (after Boeters et al., tributional patterns of a large part of the organic 1989: Fig. 149 and Shikov, 1984: 245). with world, came to an end long ago, the unraveling

of main biogeographic regions and few subdivi-

sions. What remains at a lower level, are groups of

from more or less common patterns, resulting com-

mon histories of collections of taxa that are closely

related with regard to certain biological, not neces-

sarily systematic or phylogenetic characters (migra-

tory capacities, ecological characteristics, species

longevity, etc.).

According to terrestrial snails, for example, there

is a close relation between the Greek islands of An-

tikythira and Crete, and between the island of

Kythira and the Peloponnese (Boettger, 1894; Git-

tenberger, in prep.). Comparable to Wallace's line

there is "Boettger's line" between Antikythira and

Kythira, based uponpatterns in non-marine gastro-

pods. One could expect to find Boettger's line in Fig. 2. Pleistocene (squares) and recent (dots) records of

other non-marine far these similis (Reinhardt) (after Boeters et al., 1989: Fig. 161). organisms as well, as as

have at least similar migratory capacities and about

of is the same way and speed speciation. It not sur-

birds butterflies do Phylogenetics versus area cladistics prising that for example or not

show Boettger's line; according to these organisms,

There is a fundamental difference between phylo- the southern Peloponnese, Kythira, Antikythira genetics and area cladistics. A group of species has and Crete are all equally closely related. It would be phylogenetic relationships, but at least from the absurd to ask for the "true relations" between - 153 Bijdragen tot de Dierkunde, 60 (3/4) 1990

has neither these areas. Such a "true relation" a biological nor a geological meaning.

Species dynamics and individuality

A phenomenon that has not yet received much at- tention in the biogeographical literature, at least

the in- partly for reasons just mentioned, concerns dividualistic shifts in distributionalpattern of cer- tain species during their geological history. The fact

react that the elements within an ecosystem may

climatic more or less independently to for example

for the easiness of changes, has consequences reconstructing ancient biotas. This is also relevant for our views concerning area cladistics.

Erkamo (1956), Huntley & Birks (1983) and several other authors noticedthat climatic changes did not simply induce shifts in entire vegetations, but affected the ranges of various plant species in

and records of different ways. Thus species compositions and, as Fig. 3. Miocene (squares) recent (dots)

altered fusca (Montagu) (after Boeters et al., 1989: Fig. 48). a consequence, ecological interactions, were as well, which implies much more dynamic events than would be expected at first sight. Similar obser-

of vations have been published for the ranges

and Graham species by e.g. Coope (1979)

(1986).

While revising the systematics and distributionof the terrestrial prosobranch gastropod family

Aciculidae, species of which may be found in the fossil record unchanged for several millions of

distribu- years, some additionaldata on individual tional shifts could be gathered (Boeters et al.,

1989).

Platyla polita (Hartmann, 1840) is known from

recent central Europe since the Miocene; its actual distri- Fig. 4. Miocene (square) and (dots) records of Acicula limbata Reuss (after Boeters et al., 1989: Figs. 37, 58). northern bution encompasses and, more fragmen- tary, southern Europe. In warm periods during the

the Pleistocene extends in Pleistocene the species occurred from central Eu- away from records, Italy

and southeast where it out- rope northwestward to Great Britain(Fig. 1), where Europe, occurs partly it became extinct afterwards (although it survived side the actual range of P. polita. in southern Denmark and southern Sweden). (Montagu, 1803) (Fig. 3) is known

The congeneric Platyla similis (Reinhardt, 1880) from Miocene and Pliocene deposits in central

(Fig. 2) was sympatric with P. polita during warm Europe, next to its strictly western European actual

of the Pleistocene. afterwards the Acicula limbata 1860 is also parts However, range. Reuss, (Fig. 4), former species became completely extinct in central known from Miocene deposits in centralEurope; at

site found and northernEurope. Its actual disjunct range, far one the two species are even together. 154 E. Gittenberger - Area cladistics and species individuality

However, the present range of A. limbata is in the References

Caucasus, which implies a shift opposite to that in Boettger, O., 1894. Die Binnenschnecken der griechischen In- A. fusca. seln Cerigo und Cerigotto. Nachrichtsbl. dtsch. malakozool. Pleistocene Thus the ranges of and Ges., 26 (1-2): 1-12. P. similis contribute to an area cladogram that Boeters, H.D., E. Gittenberger& P. Subai, 1989. Die Aciculidae

differs considerably from the one supported by the (: Gastropoda Prosobranchia). Zool. Verh. Leiden,

252: 1-234. recent ranges of the same species. The fossil and the Coope, G.R., 1979. Latecenozoic fossil Coleóptera: evolution, recent ranges of Acicula fusca and A. limbata do biogeography, and ecology. Ann. Rev. Ecol. Syst., 10: also illustrate this problem. Maybe we should con- 247-267.

clude that the principles of area cladistics cannot be Cracraft, J., 1988. Deep-history biogeography: retrieving the

fruitfully applied in Europe because of the environ- historical pattern of evolving continental biotas. Syst. Zool.,

37 (3): 221-236. mental changes in that part of the world, which are Erkamo, V., 1956. Untersuchungen über die pflanzenbi- not unique, however. Maybe we should neglect ologischen und einige andere Folgeerscheinungender neuzeit- of with individualistic shifts in distribu- groups taxa lichen Klimaschwankungen in Finnland. Annls. bot. Soc. tional but how do such patterns, we recognize taxa zool. bot. fenn. 'Vanamo', 28: 1-290.

if remain unknown? Should the dis- fossils we use Gittenberger,E., 1984. Vicariantists and dispersalists among the

Chondrininae (Gastropoda, Pulmonata). In: A. Solem & junct ranges of a single species in various geological

A.C. van Bruggen (eds.), World-wide snails: 56-69 (Brill, periods to derive another (? "vertical") kind of Leiden). area cladogram, highly dependent again from the Graham, R.W., 1986. Response of mammalian communities to species involved? environmental changes during the late Quaternary. In: J.M.

The present author concludes that area cladistics Diamond & T.J. Case (eds.), Community ecology: 300-313

is of a limited value. Only with regard to vast bio- (Harper & Row, New York).

Hengeveld, R., 1988. Is het heden de sleutel tot het verleden? geographic realms one could argue that these have Biovisie, 68 (7): 126-127. meaningful generalized relationships. However, Huntley, B. & H.J.B. Birks, 1983. An atlas of past and present these biogeographic relationships became known pollen maps for Europe: 0-13000 years ago: i-xiv, 1-667

even before the term area cladistics was invented. (Cambridge).

Myers, A.A. & P.S. Giller (eds.), 1988. Analytical biogeogra-

phy: i-xiii, 1-578, text-figs. (Chapman & Hall, London &

New York).

Shikov, Effects use on Acknowledgements E.V., 1984. ofland changes the land mol- lusc fauna in the central portion of the Russian plain. In: A.

Solem & A.C. van Bruggen (eds.), World-wide snails:

is cited in this short contribu- Although Hengeveld (1988) not 237-248 (Brill, Leiden). tion, his paper was a valuable introduction to the literature, which I would like to acknowledge here. Received: 20 November 1989