LANKESTERIANA 7: 33-36. 2003.

ORCHIDS SMARTER THAN SCIENTISTS – AN APPROACH TO ()

STIG DALSTRÖM

Research Associate, Marie Selby Botanical Gardens, 811 South Palm Avenue, Sarasota, FL 34236, U.S.A. [email protected]

When initiating a project in taxonomy it is Many of the color forms are probably the results of necessary to carefully examine the type specimen for cross-pollination with other species. each taxon that has been described. Just as important, however, is to explore the natural habitats of the I define a species as a group of that look plants because that is where the dynamics occur, alike regarding taxonomically important features and which is the logical source for the necessary data. It is which produce offspring with the same important fea- in nature where we best can develop an understanding tures as the parents. I distinguish a species by one or of what a “species” really is and its role in the envi- more unique important features, or a unique combina- ronment. Another fact to remember is that a previous- tion of features, which constitutes the species profile. ly described “species” does not necessarily constitute A species profile generally, but not exclusively, con- a good natural species. Specific and generic concepts sists of features that are associated with the pollina- can differ drastically between scientists even regard- tion apparatus ( and lip relation in Oncidiinae) ing the same group of plants (e.g., Dalström 2001b while a profile can consist of many other fea- versus Christenson 2002). tures as well, such as vegetative or anatomical differ- ences. A genus is a selected group of species, which It appears generally accepted that the primary basis share a unique combination of features that distin- of evolution in all organisms is mutation. Through guish them from plants in other genera. A subtribe is minor and frequent mutations an isolated population a group of selected genera, which share some unique that does not exchange genetic information with other features, or combination of features that make them populations can keep even initially insignificant different from other subtribes, etc. genetic changes inside their own group. Over many new generations of plants they will eventually have I consider the pollination of the the most sig- time to amplify the changes and become distinct. nificant part of the speciation process. Therefore, we Another possible source of new taxa is through natur- need to study the design of the pollination apparatus al hybridization where the offspring become so dras- to distinguish the various species. Some visual and tically different from the parents that they will not obvious characteristics of the flower morphology, cross-pollinate with any of them. The hybrids there- such as a larger or smaller callus on the lip, variation fore “need” to find a new or develop some in color, etc., are obvious features and may initially other isolating barrier to ensure a separate evolution- seem important. They can also be the result of ary path. It has been possible to identify many natural changes in the environment, however, such as cooler hybrids in Kunth that were originally temperatures, brighter light or dryer air, which affect described as species by crossing the suspected parent the general health of the plant. Plants in the subtribe species under artificial conditions (Rolfe 1893). Some Oncidiinae demonstrate a notorious ability to change species are more promiscuous than others and one of the flower appearance due to an environmental the worst is Lindl. Over one impact (stress). Naturally, the size and age of a plant hundred “varieties” [color forms] have been named also matters. I call this ability a plant’s natural plas- (Bockemühl 1989), which demonstrates the incredi- ticity, which is different from a plant’s natural vari- ble variability and popularity of this species alone. ability. Compared with humans one could say that 34 LANKESTERIANA Nº 7

“plasticity” represents our habit of making different species of the same superspecies. In my vocabulary, faces depending on our mood, as opposed to how dif- the term “variety” refers to individual plants that ferent individuals appear in length, nose shape or hair share some deviating features, such as a lack of pig- color, regardless of the mood (= variability). In ment, with individual plants in other populations, Odontoglossum many of the traditional distinguishing which has little taxonomic significance. features between species are “cosmetic” due to vari- Traditionally, the flower shape has dominated the ability and plasticity and do not really tell us much basis for Oncidiinae taxonomy. The angle between about the species profile. Some species descriptions the column and the lip has played a major role in how are also based on cultivated and stressed plants, we classify plants on a generic level, while features which can produce with deviating shapes. such as plant shape and habit, micro-morphology of Obviously, different species of the same genus may the pollinarium and plant anatomy have been largely also have strikingly different plant habits and anato- ignored. Vegetatively different plants have ended up my in addition to floral differences. In Odontoglos- in the same genus because they have flowers with a sum, however, the species are so vegetatively similar similar angle between the lip and the column. One of that they look pretty much the same without flowers. the problems with this approach is that the angle is They may differ in mature size and shape but a young not a definite feature, but rather a gliding scale from plant of a large species looks very much like a large the lip and column being parallel, to 180° apart, or plant of a small species. more. Both the lip and the column also show a great diversity and irregularity in shape and size, which Most species of Odontoglossum were described makes it difficult to measure the angle in a consistent during the nineteenth century by taxonomists with way. It is actually rather strange that so much atten- very little or no natural experience of the plants or tion has been paid to a few floral features when the their habitats, which led to descriptions based on very vegetative parts of the plants can show a great differ- limited knowledge. As a metaphor we can use the entiation in shape and habit. On a specific level, how- Andes hidden in mist. As the mist evaporates the ever, it seems logical to assume that the most impor- individual peaks begin to emerge. This is how orchid tant aspect of a plant’s life is the reproduction phase. species often are perceived, as isolated and distinct Therefore, the pollination strategy and dynamics must entities separated by shape, size or color. As the mist be intimately involved with a species evolution and continues to evaporate the peaks connect via interme- identity. Consequently, the taxonomically important diate ridges and eventually turn into one continuous features are those that concern the pollination appara- mountain range. Orchid species that originally appear tus (column and lip relation in Odontoglossum), distinct often become connected when more plant which need to be rather stable. material is examined from intermediate localities. Many Odontoglossum species are not distinct entities There seem to exist two antagonistic “forces” in but rather belong to a complex of similar taxa. In nature, which concern the reproduction of flowering some cases the members of a complex have satisfac- plants. One force encourages morphological stability tory profiles to justify a specific status. In other cases, to enable a species to develop some kind of consistent they connect with each other by intermediate forms. inter-action with a particular pollinator. The reason If the complex is too blurry I prefer to treat the entire why cross-pollination (the exchange of pollen complex as a superspecies, which refers to a level between two separate individual plants, or clones) is slightly “above” a more traditional species concept. A preferable to self-pollination is that the exchange of superspecies consists of subspecies, which refers to a genetic information increases variability. This, in level slightly “below” species, and are geographically turn, allows a wider adaptation to possible changes in restricted populations of plants with a shared sub-pro- the environment. If the flowers kept changing contin- file that differs from other subspecies. A subspecies is uously, however, the result would be a genetic and morphologically relatively consistent within the pop- morphological chaos, and a probable extinction due ulation and does not generally mix with other sub- to a lack of fitting . The extreme conse- Mayo 2003 DALSTRÖM - An approach to Oncidiinae taxonomy 35 quence of a too stable morphology, on the other hand, in its quest for whatever reward it is searching for. would result in an inability to adapt to changes in the Cosmetic features are here called taxonomically environment and to new pollinators, which inevitably unimportant and are unreliable for distinguishing also would lead to extinction. A balance between species. Features, such as the pollination apparatus, genetic stability and variability through mutations, that “design” the flower to accommodate particular natural hybridization and plasticity, seems to be the pollinators are taxonomically important features. successful choice. It is simple to discern a successful These cannot change too much because it would lead strategy versus an unsuccessful one in Odontoglos- to a morphological chaos with consequent pollination sum because the plants either exist commonly or not, problems. Unfortunately, what works to fool pollina- and the majority of Odontoglossum species display a tors also fools scientists. Only by visiting many popu- great variability. It seems logical that plants with a lations and studying large quantities of plants can we variable gene-code have a higher survival rate in an see through this deception and discern natural unstable environment, than do plants with a more species. fixed genetic constitution. The Andean region of When we fully understand the evolutionary dynam- South America provides a constantly changing envi- ics in a small group of plants, such as ronment due to volcanic eruptions, earthquakes, land- Odontoglossum, we discover a valuable key to the slides and floods, and more recently, man’s occur- understanding of how other organisms evolve on rence on the stage. In a longer perspective, fluctuation Earth as well. Repetitive speciation patterns on many in temperature has caused periods of glaciations as levels in plants have lead to the development of well. Consequently, a variable habitat would also superficially similar flowers based on similar pollina- favor a rapid and diverse speciation, which seems to tion strategies. Using the flower morphology exclu- be the case in the Andes. sively for taxonomic treatments may seem convenient The flowers of Odontoglossum are considered and user-friendly but can be misleading and confus- rewardless. They do not offer any kind of obvious ing at the same time because it does not necessarily reward, such as nectar or oil for the pollinator. relate to the natural relationship between species and Theoretically, a pollinator may visit a seemingly genera. In contrast, with the recent development of rewardless flower for some other rewarding reasons, DNA sequence analysis we have found another key to such as absorbing a scent or possibly to deposit eggs the classification of troublesome plants. Although on the flower, which in turn may feed the larvae. No still at an experimental, initial stage of its develop- such observations are documented, however. In order ment, it is now possible to learn about the genetic to encourage the pollinator to re-visit a rewardless relationships of the plants. This can be very useful in flower the plant has to develop a deceit pollination some cases, such as the delineation and separation of strategy, and this seems to be done with the help of Cyrtochilum from allied genera (Dalström 2001a). variable and plastic features, or cosmetics. Examples Unfortunately it also creates a new dimension of of cosmetics are coloration patterns, variation in the problems when we realize that morphologically dif- callus size and shape, width of and , and ferent looking plants can be closely related, and possibly a variation in odors (e.g. the different parts viceversa, thus breaking up traditional classification of the flower of Lindl. have dif- and creating a very unpractical system. In other ferent smells). A certain degree of natural hybridiza- words, with a classification system based solely on tion also seems favorable to an increased variability DNA analysis, you may have to know the plant before you can identify it, because there are no reli- as long as the pollination strategy is not broken up able and visible indications where it belongs. and becomes chaotic. In conclusion, we have certain features, which vary not only between different indi- LITERATURE CITED vidual plants but also between individual flowers on Bockemühl, L. 1989. Odontoglossum, a Monograph and the same and are assumed to be deceptive. Iconograph. Brücke-Verlag Kurt Schmersow, They convince the pollinator to make yet another try Hildesheim. Germany. 36 LANKESTERIANA Nº 7

Christenson, E. A. 2002. Cochlioda - A taxonomic treat- Dalström, S. 2001b. New species and combinations in the ment of this New World genus. Orchids 2(110–121). Oncidiinae (Orchidaceae) and a synopsis of the Dalström, S. 2001a. A synopsis of the genus Cyrtochilum Cochlioda clade. Selbyana 22(2): 135–145. (Orchidaceae; Oncidiinae): Taxonomic reevaluation and Rolfe, R. A. 1893. Hybrid . Orch. Rev. new combinations. Lindleyana 16(2): 56–80. 1(5): 142.

Stig Dalström is a free-lance botanical artist, and Research Associate of the Marie Selby Botanical Gardens, Sarasota, Florida, US. He specializes in Oncidiinae taxonomy, particularly the Andean species of Cochlioda, Cyrtochilum, and Odontoglosum.