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Analysis of a Species/Instars/Characters Table: a Theoretical Survey Contributions to Zoology, 67 (3) 197-220 (1998) SPB Academic Publishing bv, The Hague of theoretical Analysis a species/instars/characters table: a survey on the use of chaetotaxy in ontophylogenetic studies Pierre Nayrolles Laboratoire de Zoologie, Ecobiologie des Arthropodes édaphiques, Université Paul Sabatier, 118 route de Narbonne F-31062 Toulouse Cedex France. E-mail: , [email protected] Keywords: Ontogeny, phytogeny, chaetotaxy, probabilistic organs, ontogenetic trajectories, arthropods, Collembola, Symphypleona Abstract opposant présences et absences. Dans une population, une soie donnée peut se révéler variable, une probabilité de présence studies of Phylogenetic on several groups arthropods, such as pouvant alors lui être attachée. Une soie variable pourrait être Acari (mites) or Collembola (springtails), make wide use of al- perçue comme la marque d’un polymorphisme (plusieurs chaetotaxy. Chaetotaxic characters, besides being morpho- lèles dans une population). En fait, cette variabilité doit être logical features (setal shape), often have a binary nature, that comprise comme la sommation de propensions individuelles, A be variable is presence vs. absence. seta can in a popula- ou en d’autres termes, comme la résultante d’une potentialité and to this seta. tion, one may attribute a presence probability à individu hasard. propre chaque exprimée au Vraisembla- In the for fact, presence probability should be defined each blement, un phénomène similaire à celui de l’inhibition laté- instar. One might think that a variable seta corresponds to a rale, phénomène qui explique comment une fluctuation au polymorphism (e.g., two or more alleles in a population); in hasard intervenue précocement peut déterminer le destin de of fact, setal variability should be regarded as the result a à des Un cellules, est l’origine organes probabilistes. tableau propensity intrinsic to individuals, i.e., a potentiality being tridimensionnel espèces/stades/caractères contenant les pro- random known expressed at among specimens. The phenomenon babilités de présence des soies peut être constitué. Deux sortes as lateral inhibition explains how an early random fluctuation de traitements, dans lesquels le problème des soies homéo- is of at the root the cell fate. Probabilistic organs are likely to types prend une place essentielle, sont alors applicables. Les from similar originate a phenomenon. A tridimensional spe- autres questions relatives à ces analyses sont la pertinence cies/instars/characters table (SIC table), containing setal pres- de l’utilisation des probabilités de soies et la réduction de be Two kinds of ence probabilities, can built up. analyses l’information redondante. Un premier traitement conduit à un may be applied to such a table. In these analyses, the problem tableau bidimensionnel dont les caractères sont en colonnes of homeotypic setae is emphasized. Related issues are the divisées et, en lignes, les espèces en stades. En utilisant des of setal and the of accuracy using probabilities problem re- statistiques multidimensionnelles, il est possible de dessiner of information. A first ducing redundancy process leads to a des et donc de les trajectoires ontogénétiques, comparer on- bidimensional in table with characters columns, and the spe- togenèses. Le but du deuxième traitement est l’obtention d’un cies divided into their instars in rows. By using multidimen- tableau être utilisé dans espèces/caractères qui pourra une sional statistics we can access ontogenetic trajectories, and in analyse cladistique. this way, ontogenetic comparisons can be achieved. The aim of the second process is to produce a species/characters table which can be used in a cladistic analysis. Introduction Résumé that several authors Andre (1988) emphasized were surprised that so little is known about insect im- Dans l’étude de phylogénétique plusieurs groupes d’Arthro- mature stages. He explained this fact by the lack tels les podes, que Acariens ou les Collemboles, il est large- of a method for the treatmentof ontogenetic data. ment fait appel à la chétotaxie. Outre le fait qu’ils puissent The aim of this as well as those Andre, être des traits purement morphologiques (forme des soies), paper, by les caractères chétotaxiques sont souvent de nature binaire, is to provide some theoretical ideas and several Downloaded from Brill.com10/10/2021 04:06:52PM via free access 198 P. Nayrolles - Chaetotaxy in ontophylogenetic studies techniques for studying arthropod ontophyloge- merit of mites and distinct from each other by “all netics (i.e., ontogenetics and phylogenetics con- or none” characters. sidered together) by means of chaetotaxy, i.e., The discrete characters used for differentiating the of the of - based the of study arrangement organs setae or, stases are generally on presence se- - considered this strictly speaking, sensilla produced by the in- tae. The characters in paper con- tegument of arthropods. Once chaetotaxic pat- cern only idionymic setae. Idionymy (Grandjean, have absence is the of terns been established, presence vs. 1949) “quality” a particular organ as or number of setae can be used to compare spe- distinct from other organs of the same nature; defined cies. such an organ occupies a position so that A general scientific procedure by which char- it can be designated by a name or a symbol, e.g., acters would be generated cannot be defined, be- our vertebrae are idionymic while our hairs are cause of the great diversity of features used (e.g., not. Idionymy of a set of setae is established by morphological, behavioral, biochemical features, comparisons of specimens within a species. De- etc.). Nevertheless, in some fields, a standard velopmental stage or sex should also be consid- method be set ered because setae be in larvae can up (e.g., morphometry). may idionymic This article focuses on the erection of charac- and not in adults. When setae are not idionymic, ters from brute chaetotaxic data with a special their positions are not constant but depend on attention paid to the problem of variable setae. their number, in other words the inter-setal dis- is of the setal number. Ontogeny, i.e., instar of appearance of setae, is tance a function Converse- used that char- when each has as a way to distinguish species, so ly, setae are idionymic, seta a fixed and if is the acters should embrace the ontogenetic dimension. position, a seta absent, position Polarization of characters and phylogenetic in- of the neighboring setae is not modified. ference and will discussed Setae that named are next steps not be appear during development are while setae” here. Even if phylogenetics is not directly ad- “secondary setae”, “primary are from dressed, this theme is underlying since building present the first instar on. Sometimes, at a up characters is a preliminary phase before any given instar, some setae are variable. Setal pres- phylogenetic - cladistic - work. ence probabilities must be defined at the level of An of is taken from In this the of example data processing a population. paper, concept proba- the group of Symphypleona, one of the major bilistic organs will be emphasized. divisions of the Collembola (Hexapoda). The onto- Analysis of a SIC table permits ontophyloge- genetic patterns of these arthropods are quite simple netic studies. For instance, it is possible to carry and constitute for the method out multidimensional statistics shed a good support pre- to light on sented. similarities dissimilarities instars of or among spe- The of cies. A dealt with of these postembryonic development arthropods recent paper application consists of a succession of forms separated by statistical methods (Nayrolles, 1996), but missing of life between molts the from it discussion the and molts. The period two is was a of nature possi- “intermolt”, also called “instar”. This paper de- ble uses of chaetotaxic characters. From my study of scribes a method for analyzing a table with three Symphypleona, it appears that data processing dimensions, these dimensions being species, their of chaetotaxic characters is rather burdensome instars, and characters, in other words a species/ because of the numerous setae that have to be instars/characters (SIC) table. In principle, my taken into account. Computer software wouldmake method can be applied to all arthropod species studies of SIC tables easier, but before undertak- exhibiting “stases” and associated discontinuous ing such a development it is important to outline The word the these characters. “stase” was coined by Grand- purpose and methods of analyses as for the and jean (1938) designating morphological stag- well as to specify properties of a SIC table its es succeeding one another during the develop- subsets. Downloaded from Brill.com10/10/2021 04:06:52PM via free access 67 - Contributions to Zoology, (3) 1998 199 Three basic and concepts: species, instar, to simplify the situation, sex will be left out to character avoid having four dimensions in the SIC table. Nixon & Wheeler (1990a, 1990b) made a dis- Andre (1986, discussed the of 1988) concept stase tinction between attributes that and vary attributes and proposed to extend it to other arthropods, that are constantly distributed among specimens particularly Collembola. According to Andre of an elementary division of the species-instar- each (1989a), intermolt of a mite or Col- juvenile sex The first spectrum. type was named a trait, lembola is a and the adult of stase, Collembola, the second a character, or more precisely, a char- which molts but does discontinu- not change by Three acter state. categories were established: 1) also ous characters, corresponds to one stase. Al- the trait as a variable attribute fitting with varia- the of and instar are differ- though concepts stase the bly distributed alleles, 2) character state as ent, I will use the word “instar” instead of “stase”, constant attribute permitting species diagnoses, because it has the of better known and advantage being 3) the character as a set of character states is based on among entomologists. My argument derived from one another through a series of trans- of a but Andre’s commonness term, conceptually formations.
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