全 文 ：Journal of Systematics and Evolution 46 (3): 396–404 (2008) doi: 10.3724/SP.J.1002.2008.08039
(formerly Acta Phytotaxonomica Sinica) http://www.plantsystematics.com
Evolution of fruit types and seed dispersal:
A phylogenetic and ecological snapshot
Claire M. LORTS Trevor BRIGGEMAN Tao SANG*
(Department of Plant Biology, Michigan State University, East Lansing, MI 48824, USA)
Abstract Success of flowering plants is greatly dependent on effective seed dispersal. Specific fruit types aid
different mechanisms of seed dispersal. However, little is known about what evolutionary forces have driven the
diversification of fruit types and whether there were phylogenetic constraints on fruit evolution among angio-
sperm lineages. To address these questions, we first surveyed the orders and families of angiosperms for fruit
types and found no clear association between fruit types and major angiosperm lineages, suggesting there was
little phylogenetic constraint on fruit evolution at this level. We then surveyed fruit types found in two contrasting
habitats: an open habitat including the Indian desert and North American plains and prairies, and a closed forest
habitat of Australian tropical forest. The majority of genera in the survey of tropical forests in Australia were
fleshy fruit trees, whereas the majority of genera in the survey of prairies and plains in central North America
were herbs with capsules and achenes. Both capsules and achenes are frequently dispersed by wind in the open,
arid habitat, whereas fleshy fruits are generally dispersed by animals. Since desert and plains tend to provide
continuous wind to aid dispersal and there are more abundant mammal and bird dispersers in the closed forest,
this survey suggests that fruit evolution was driven at least in part by dispersal agents abundant in particular
habitats.
Key words adaptation, angiosperm, animal dispersal, fruit development, wind dispersal.
Seed dispersal is essential for the success of plant
reproduction and adaptation. Germination and growth
away from the mother plant allows opportunities to
find advantageous areas to inhabit, in addition to
avoiding unfavorable conditions around the mother
plant such as inbreeding and sibling competition
(Willson & Traveset, 2000). The origin of fruits of
angiosperms was a major evolutionary innovation that
greatly enhanced seed dispersal efficiency and trig-
gered rapid diversification of flowering plants. How-
ever, little is known about the evolutionary force that
drove rapid diversification of fruit types.
Fossil records since the Cretaceous show a diver-
sity of dispersal mechanisms present in both gymno-
sperms and angiosperms. Abiotic seed dispersal
(mostly wind dispersal) was most prominent during
the Cretaceous period, but a small portion of possible
animal-dispersed seeds may have existed in both
gymnosperms and angiosperms (Eriksson et al.,
2000a). Although animal dispersal existed in gymno-
sperms and angiosperms before the Tertiary, dinosaurs
were not frugivores and closed forest systems were
not present, therefore providing little selection pres-
sure on earlier production of fleshy fruits (Tiffney,
2004). At the beginning of the Tertiary period, several
events might have had substantial impact on fruit
evolution, especially the marked increase in the
abundance and diversity of fleshy fruits (Eriksson et
al., 2000a, b; Tiffney, 2004).
One event was climatic change, featured by
global temperature cooling down and an increase in
moisture. The other was the extinction of dinosaurs
and subsequent radiation of birds and mammals. Both
events allowed the development of closed forest
systems in the early Tertiary and the radiation of
fleshy fruits, which was coupled with animal diversi-
fication in the forest habitats (Bolmgren & Eriksson,
2005).
The development of closed forests had several
consequences. First, little wind travels through closed
systems, making wind dispersal, especially for under-
story plants, more difficult. Second, the lack of light
in closed forest communities favored larger seeds
whose seedlings could have better chances to establish
under the shaded conditions. The large seeds were
more effectively dispersed by animals than by wind.
Additionally, animals tended to deposit seeds in
canopy gaps where there was adequate light to facili-
tate seedling establishment (Bolmgren & Eriksson,
2005). Finally, the closed forest systems increased the
chance of interaction between plants and birds and
mammals. The interaction was mutually beneficial,
with plants providing nutritious fleshy fruits to
———————————
Received: 26 March 2008 Accepted: 23 April 2008
* Author for correspondence. E-mail: .
LORTS et al.: Fruit evolution

397
animals and animals serving as effective agents for
seed dispersal (Herrera, 1989; Clark et al., 2001).
These factors greatly facilitated the co-radiation of
fleshy fruits of angiosperms and the species of birds
and mammals in the early Tertiary.
To better understand the evolution of fruit types
during angiosperm diversification, we took two
snapshots of the problem from phylogenetic and
ecological perspectives. At first, we surveyed fruit
types of all angiosperm orders and mapped them onto
the angiosperm phylogeny. In this survey, we aimed
to test the hypothesis that there were major shifts in
fruit types during angiosperm diversification. If this
was true, we should be able to observe shifts in fruit
types associated with branches of the angiosperm
phylogeny.
We surveyed 59 orders and 391 families for
available fruit types (Table 1) from numerous web-
sites and books (e.g., Angiosperm Phylogeny Website
http://www.mobot.org/MOBOT/Research/APweb/wel
come.html; The Families of Flowering Plants http://
delta-intkey.com/angio/; Heywood, 1993; Kubitzki et
al., 1993; Judd et al., 2002). We grouped various fruit
types into three major categories: fleshy, dry dehis-
cent, and dry indehiscent. Fleshy fruits included
berries, drupes, and pomes. Dry dehiscent fruits
consisted of follicles and capsules (including siliques).
Dry indehiscent fruits included samaras, schizocarps,
nuts, and achenes (including utricles and caryopsis).
The fruit types were mapped onto the phylogenic tree
from the Angiosperm Phylogeny Website. The visual
inspection did not detect any association between fruit
types and major clades (Fig. 1). Multiple fruit types
are scattered across the angiosperm phylogeny. This
indicates that there was little phylogenetic constraint
on fruit evolution at least in this evolutionary scale.
Fruit types may have evolved repeatedly within orders
under natural selection.
This led us ask the question of whether fruit
evolution has been driven by dispersal mechanisms. If
the answer is positive, fruit types favored by dispersal
mechanisms prevalent in certain habitats should be
more abundant than other fruit types. To test this
hypothesis, we conducted a second survey looking at
fruit types present in different types of habitats. We
predicted that fleshy fruits would be abundant in
dense forest habitats that inhabit many vertebrates and
birds, whereas open habitats with more available wind
would have a higher proportion of dry fruits fre-
quently dispersed by wind.
In addition to habitats, we considered that growth
form was also important because different growth
forms have advantages for seed dispersal in certain
habitats more than others. Willson and Traveset
(2000) suggested that wind dispersed plants were tall
relative to other plants in the habitat. Taller trees in
the forests have better access to wind currents and
therefore the seeds are more likely to be dispersed by
wind. In contrast, understory plants in closed forest
systems tend to not have access to enough wind for
efficient dispersal, and may consequently favor other
dispersal mechanisms.
We surveyed two contrasting habitats, with
Australian tropical forests representing the closed
forest system and both Central North American plains
and Indian desert representing open habitats. The total
of 786, 922, and 247 genera were surveyed for the
Australian tropical forests (Cooper, 2004), Central
North American plains (Rydberg, 1932), and Indian
desert (Bhandari, 1978), respectively (The data sets
are available from the authors upon request). We
compared animal-dispersed fleshy fruits and fruits
favored by wind dispersal including samara, capsule,
and achene (Pijl, 1982). Follicles and nuts were
treated as a separate category because dispersal
mechanism varies between abiotic and biotic in these
two fruit types. Growth forms were grouped into trees,
shrubs to small trees, herbs, and vines or mistletoes.
Several genera had more than one fruit type and/or
growth form, and in some cases a variant fruit type
matched up with a specific variant growth form. To
account for these variations we made an additional
value for a unique variation in a genus. For instance,
when a genus had only one growth form and two
different fruit types, 2 values were used to separate the
fruit types. Some genera had two growth forms and
two fruit types, in which case two separate values
were assigned, but the specific fruit type and growth
form it corresponded to were coupled.
The results of the survey are presented as percent
of genera that had particular fruit types in the given
habitat. Figures 2A, C, and E display values for fruit
types of each growth form and Figures 2B, D, and F
help visualize the relative abundance of fruit types in
each habitat. The fruit types are categorized according
to the most frequent dispersal mechanism in the
habitats, including fleshy fruits primarily dispersal by
animals, dry fruits (capsule, samara, achene) often
dispersed by wind, and other dry fruits (follicle and
nut) with uncertain frequency of biotic and abiotic
dispersal.
Australian tropical forests had the most diverse
fruit types and growth forms (Fig. 2: A, B). The ma-
jority of genera were fleshy fruit trees. The following

Journal of Systematics and Evolution Vol. 46 No. 3 2008 398
Table 1 Fruit types surveyed in families of angiosperms. Families were grouped into orders corresponding to the phylogenic tree from the
Anigosperm Phylogeny Website
Order Family Fruit type Order Family Fruit type
Amborellales Amborellaceae drupe Luzuriagaceae capsule, berry
Nymphaceae Cabombaceae follicle, achene Alstroemeriaceae capsule, berry
Nymphaeaceae berry, nut Rhipogonaceae berry
Austrobaileyales Austrobaileyaceae berrylet Philesiaceae berry
Illiciaceae Follicle Smilacaceae berry
Trimeniaceae berry Liliaceae capsule, berry
Chloranthales Chloranthaceae drupaceous Asparagales Orchidaceae capsule
Magnoliales Myristicaceae follicle (aril) Blandfordiaceae capsule
Magnoliaceae follicle (aril), samara Lanariaceae capsule
Degeneriaceae follicle Asteliaceae berry, capsule
Himantandraceae drupe Hypoxidaceae berry, capsule
Annonaceae berry Ixioliriaceae capsule
Laurales Calycanthaceae achene, drupe Tecophilaeaceae capsule
Gomortegaceae drupe Doryanthaceae capsule
Atherospermataceae achene Iridaceae capsule
Monimiaceae drupe, achene Xanthorrhoeaceae capsule
Hernandiaceae samara, nut Hemerocallidaceae capsule, berry, nut
Lauraceae drupe, berry Asphodelaceae capsule
Canellales Winteraceae berry, follicle Alliaceae capsule
Canellaceae berry Amaryllidaceae capsule, berry
Piperales Aristolochiaceae capsule, berry, follicle Aphyllanthaceae capsule
Piperaceae drupe Hyacinthaceae capsule
Saururaceae capsule, or dry indehiscent Agavaceae capsule
Acorales Acoraceae berry Asparagaceae berry
Alismatales Araceae berry, capsule Ruscaceae berry
Hydrocharitaceae berry, capsule Unplaced Dasypogonaceae capsule
Alismataceae achene Arecales Arecaceae berry, drupe
Limnocharitaceae follicle Poales Rapateaceae capsule
Scheuchzeriaceae follicle Sparganiaceae drupe, nut
Aponogetonaceae follicle Typhaceae drupe, achene like follicle
Juncaginaceae achene, follicle Bromeliaceae capsule, berry
Posidoniaceae spongy pericarp,dehiscent Thurniaceae capsule
Ruppiaceae drupelet Juncaceae capsule
Cymodoceaceae achene, drupelet Cyperaceae achene
Zosteraceae achene Mayacaceae capsule
Potamogetonaceae achene, drupe Eriocaulaceae capsule
Petrosaviales Petrosaviaceae follicle Xyridaceae capsule
Dioscoreales Nartheciaceae capsule Anarthriaceae nut
Taccaceae berry, capsule Centrolepidaceae follicle
Thismiaceae capsule Restionaceae capsule, achene, nut
Burmanniaceae capsule Flagellariaceae drupe
Dioscoreaceae capsule, berry, samara Joinvilleaceae drupe
Pandanales Velloziaceae capsule Ecdeiocoleaceae achene, capsule
Triuridaceae achene Poaceae achene
Stemonaceae capsule (indehiscent of dehiscent) Commelinales Commelinaceae capsule
Pandanaceae drupe, berry Hanguanaceae drupe
Cyclanthaceae berry Philydraceae capsule
Liliales Corsiaceae capsule Haemodoraceae capsule
Campynemataceae capsule Pontederiaceae capsule, nut
Melanthiaceae capsule Zingiberales Musaceae berry
Petermanniaceae berry Heliconiaceae drupe, schizocarp
Colchicaceae capsule Strelitziaceae capsule (with aril)

LORTS et al.: Fruit evolution

399
Table 1 (continued)
Order Family Fruit type Order Family Fruit type
Lowiaceae capsule Cactaceae berry
Cannaceae capsule Santalales Erythropalaceae drupe
Marantaceae capsule, berry Olacaceae drupe, nut
Zingiberaceae capsule, berry Misodendraceae achene, nut
Costaceae capsule, nut, achene Loranthaceae berry, samara, drupe, nut
Ranunculales Eupteleaceae samara Opiliaceae drupe
Lardizabalaceae follicle, berry Balanophoraceae drupe, nut
Circaeasteraceae achene Santalaceae drupe, nut
Menispermaceae drupe Viscaceae berry
Berberidaceae berry Saxifragales Cercidiphyllaceae capsule, follicle
Ranunculaceae follicle, achene, berry Peridiscaceae drupe, capsule
Papaveraceae capsule Daphniphyllaceae drupe
Sabiales Sabiaceae drupe Hamamelidaceae capsule
Proteales Nelumbonaceae nut Altingiaceae capsule
Platanaceae achene Paeoniaceae follicle
Proteaceae follicle, nut, achene, drupe Crassulaceae follicle
Trochodendrales Trochodendraceae follicle Tetracarpaeaceae follicle
Buxales Didymelaceae drupelet Penthoraceae follicle
Buxaceae capsule, drupe Haloragaceae drupe, nut
Gunnerales Gunneraceae drupe, nut Cynomoriaceae nut
Myrothamnaceae follicle Iteaceae capsule
Berberidopsidales Aextoxicaceae drupe (dry) Pterostemonaceae capsule
Berberidopsidaceae berry Grossulariaceae berry
Dilleniales Dilleniaceae follicle, achene Saxifragaceae capsule
Caryophyllales Rhabdodendraceae drupelet Vitales Vitaceae berry
Droseraceae capsule Zygophyllales Krameriaceae achene
Nepenthaceae capsule Zygophyllaceae capsule
Drosophyllaceae capsule Celastrales Celastraceae capsule, drupe
Ancistrocladaceae nut Lepidobotryaceae capsule
Dioncophyllaceae capsule Parnassiaceae capsule
Frankeniaceae capsule Pottingeriaceae capsule
Tamaricaceae capsule Oxalidales Connaraceae follicle, drupe
Plumbaginaceae capsule Oxalidaceae capsule, berry
Polygonaceae nut, achene Cunoniaceae follicle, drupe, nut, capsule
Simmondsiaceae capsule Brunelliaceae follicle
Asteropeiaceae capsule Cephalotaceae follicle
Physenaceae capsule Elaeocarpaceae capsule, drupe, berry
Caryophyllaceae capsule, nut Malpighiales Achariaceae berry
Achatocarpaceae berry (with aril) Violaceae capsule, berry
Amaranthaceae achene, capsule Salicaceae capsule, berry, drupe
Stegnospermataceae capsule Lacistemataceae capsule
Limeaceae capsule Turneraceae capsule
Lophiocarpaceae achene, capsule Malesherbiaceae capsule
Barbeuiaceae capsule Passifloraceae berry, capsule
Aizoaceae capsule, berry, nut Goupiaceae drupe
Phytolaccaceae berry Putranjivaceae drupe
Nyctaginaceae achene, nut Lophopyxidaceae samara
Molluginaceae capsule Clusiaceae capsule, berry, drupe
Halophytaceae nutlet Bonnetiaceae capsule
Basellaceae utricle Hypericaceae berry, drupe
Montiaceae capsule Malpighiaceae samara, drupe, berry, nut
Didiereaceae capsule Elatinaceae capsule
Portulacaceae capsule Ochnaceae drupe, berry

Journal of Systematics and Evolution Vol. 46 No. 3 2008 400
Table 1 (continued)
Order Family Fruit type Order Family Fruit type
Medusagynaceae capsule, follicle Sapindaceae capsule, drupe, berry
Quiinaceae berry, follicle Nitrariaceae capsule, nut
Phyllanthaceae drupe Huerteales Dipentodontaceae capsule
Picrodendraceae drupe Gerrardinaceae berry
Balanopaceae drupe Tapisciaceae drupe, berry
Trigoniaceae capsule, samara Malvales Neuradaceae capsule, follicle
Dichapetalaceae drupe Thymelaeaceae berry, drupe, achene-like
Chrysobalanaceae drupe Sphaerosepalaceae capsule
Euphroniaceae capsule Bixaceae capsule
Caryocaraceae drupe Cistaceae capsule
Centroplacaceae capsule Sarcolaenaceae achene, nut, capsule
Ctenolophonaceae nut Dipterocarpaceae nut
Erythroxylaceae drupe Muntingiaceae berry
Humiriaceae drupe Malvaceae capsule, nut, follicle
Irvingiaceae drupe, samara Brassicales Akaniaceae capsule
Ixonanthaceae capsule Tropaeolaceae samara, nut, drupe
Linaceae capsule, drupe, nut Moringaceae capsule
Pandaceae drupe, capsule Caricaceae berry
Rafflesiaceae berry, capsule Setchellanthaceae capsule, silique
Rhizophoraceae capsule, berry Limnanthaceae nut
Cucurbitales Anisophylleaceae drupe, samara, capsule Koeberliniaceae berry
Corynocarpaceae drupe Bataceae drupe
Coriariaceae achene, nut Salvadoraceae berry, drupe
Cucurbitaceae berry, capsule Pentadiplandraceae berry
Tetramelaceae capsule Resedaceae berry, capsule, follicle
Datiscaceae capsule Gyrostemonaceae achene
Begoniaceae capsule, berry Tovariaceae berry
Fagales Fagaceae capsule, nut Capparidaceae berry
Myricaceae drupe, achene Cleomaceae silique
Juglandaceae nut, drupe, nutlet Brassicaceae berry, capsule, silique
Rhoipteleaceae samara Cornales Loasaceae capsule
Ticodendraceae drupe Hydrangeaceae drupe, samara
Betulaceae achene, samara, nut Nyssaceae drupe
Casuarinaceae samara Cornaceae drupe
Fabales Polygalaceae capsule, samara, drupe, berry, nut Curtisiaceae drupe, achene-like
Surianaceae berry, drupe, nut Grubbiaceae capsule
Fabaceae samara, follicle, achene, drupe, berry Hydrostachyaceae capsule, drupe
Rosales Barbeyaceae nut Ericales Balsaminaceae capsule
Cannabaceae achene, drupe Marcgraviaceae berry
Dirachmaceae follicle Tetrameristaceae capsule
Elaeagnaceae achene (drupe-like) Polemoniaceae capsule
Moraceae drupe, achene Fouquieriaceae capsule
Rhamnaceae drupe, samara Sladeniaceae capsule
Rosaceae achene, drupe, follicle, pome Pentaphylacaceae berry
Ulmaceae nut, samara Sapotaceae berry
Urticaceae drupe, achene Ebenaceae drupe
Celtidaceae drupe Maesaceae berry, drupe
Sapindales Kirkiaceae achene, nut Theophrastaceae capsule
Rutaceae drupe, berry, samara, follicle, capsule Primulaceae drupe
Meliaceae samara, drupe Myrsinaceae berry, capsule
Simaroubaceae drupe Mitrastemonaceae capsule
Anacardiaceae drupe, dehiscent drupe Theaceae drupe, berry
Burseraceae capsule, berry, drupe, samara Symplocaceae capsule, drupe, samara

LORTS et al.: Fruit evolution

401
Table 1 (continued)
Order Family Fruit type Order Family Fruit type
Styracaceae capsule Verbenaceae capsule
Diapensiaceae capsule, berry Boraginaceae drupe, samara
Actinidiaceae capsule Solanales Montiniaceae capsule
Roridulaceae capsule Sphenocleaceae capsule
Sarraceniaceae capsule Hydroleaceae capsule
Clethraceae drupe, samara Convolvulaceae berry, capsule
Cyrillaceae capsule, berry, drupe Solanaceae capsule, drupe, nut
Ericaceae capsule Aquifoliales Cardiopteridaceae drupe
unplaced Oncothecaceae drupe Stemonuraceae berry
Icacinaceae samara Phyllonomaceae drupe
Garryales Eucommiaceae berry Helwingiaceae drupe
Garryaceae nut, drupe Aquifoliaceae drupe
unplaced Boraginaceae capsule, drupe, berry unplaced Polyosmaceae capsule, berry
Gentianales Rubiaceae capsule Escalloniaceae follicle, capsule
Gentianaceae capsule, drupe, berry, follicle Paracryphiaceae capsule
Loganiaceae capsule Sphenostemonaceae drupe
Gelsemiaceae follicle, drupe, berry Asterales Rousseaceae capsule, berry
Apocynaceae capsule Campanulaceae berry
Lamiales Plocospermataceae capsule, samara, drupe, berry Pentaphragmataceae berry
Oleaceae nut Alseuosmiaceae drupe
Tetrachondraceae capsule Phellinaceae capsule, drupe
Calceolariaceae capsule, berry Argophyllaceae capsule
Gesneriaceae capsule Stylidiaceae capsule, berry
Acanthaceae capsule, berry Menyanthaceae capsule, drupe, nut
Bignoniaceae capsule Goodeniaceae achene-like
Byblidaceae capsule Calyceraceae achene
Cyclocheilaceae drupe, nutlet Asteraceae drupe
Lamiaceae capsule Dipsacales Diervillaceae drupe, berry, capsule, achene
Lentibulariaceae capsule Caprifoliaceae berry, drupe
Martyniaceae schizo (drupelet), drupe Linnaeaceae achene-like
Myoporaceae capsule Morinaceae achene-like
Paulowniaceae schizo (nut), capsule Dipsacaceae samara, achene
Phrymaceae capsule Apiales Torricelliaceae berry
Plantaginaceae berry Griseliniaceae drupe, berry
Schlegeliaceae capsule, achene Araliaceae capsule, berry
Scrophulariaceae capsule Pittosporaceae samara
Stilbaceae drupe, nutlet Myodocarpaceae drupe



most abundant plants were vines and shrubs/small
trees with fleshy fruits, and trees and shrubs/small
trees with capsules. It should be noted that about one
fifth of genera with capsules surveyed in this habitat
had arils, and the majority of them were trees. This is
worthy of attention since fruits with arils are com-
monly bird dispersed. Genera with samaras or achenes
were less abundant.
Most angiosperm genera surveyed in the Prairies
and Plains of Central North America were shrubs and
herbs (Fig. 2: C, D). A high proportion of genera were
herbs with capsules, followed by herbs with achenes.
These two fruit types are generally dispersed by wind
in this habitat. The very few fleshy genera had various
growth forms. Similarly, the majority of genera in the
survey of the Indian desert are herbs with capsules and
achenes (Fig. 2: E, F). Shrubs/small trees had the
highest proportion of fleshy fruits relative to other
growth forms in the Indian desert, with about an equal
ratio of fleshy to wind dispersed fruits. Desert trees
produced a higher ratio of wind dispersed fruits.
Overall, the majority of genera in the survey of
tropical forests in Australia were fleshy fruit trees,
while the majority of genera in the survey of prairies
and plains in central North America were herbs with
capsules and achenes. Both capsules and achenes are
Journal of Systematics and Evolution Vol. 46 No. 3 2008 402



Fig. 1. Fruit types mapped on the phylogeny of angiosperm orders. Categories of fruit types include fleshy fruits (white bars), dry dehiscent fruits
(back bars), and dry indehiscent fruits (hatched bars).


frequently dispersed by wind in these open and arid
habitats, while fleshy fruits are generally dispersed by
animals. Since desert and plains tended to provide
continuous wind to aid dispersal, there may not be
selective pressures for producing fleshy fruits that are
developmentally more costly. However, the high
abundance of fleshy fruits in the survey of Australian
tropical forests may indicate a possible clue to selec-
tion pressures on angiosperms for fleshy fruits due to
the high proportion of available mammal and bird
dispersers relative to the open habitats. Moreover,
trees, shrubs, and vines that have the positional ad-
vantage to attract birds have a higher proportion of
fleshy fruits than herbs. Therefore, the survey supports
the hypothesis that fruit evolution was driven at least
in part by dispersal agents abundant in particular
habitats.
This study perhaps raises more questions than it
answers. While we showed that there lacked a phy-
logenetic constraint on fruit type evolution across
major lineages of angiosperms, one could easily ask
how far down the phylogenetic hierarchy this holds
true. There have been at least several molecular
phylogenetic studies showing that the same fruit type
evolved independently within an angiosperm family
(Morgan et al., 1994; Smith, 2000; Knapp, 2002;
Zjhra et al., 2004; Motley et al., 2005), suggesting that
the phylogenetic constraint remains weak for families
with variable fruit types. Many more investigations
are needed at the familial and lower levels before any
generalization can be made.
The ecological survey also needs to be expanded
in habitat types and sample sizes so that the hypothe-
sis can be statistically tested. Whenever possible,
empirical observation should be conducted to conform
the dispersal mechanisms, especially for fruits with
unclear or multiple dispersal agents. For instance, in
this survey capsules (not including capsules with arils)
were considered to be generally wind dispersed.
However, some plants with capsules have ballistic (or
LORTS et al.: Fruit evolution

403




Fig. 2. Survey of fruit types in different habitats. A, C, E, Percent abundance of fruit types of specific growth forms in the Australian Tropical
Forest, Prairies and Plains of Central North America, and Indian Desert, respectively. White bars: fleshy fruits; black bars: dry fruits from left to right,
capsule, samara, and achene; hatched bars: dry fruits from left to right, follicle and nut. B, D, F, Percent abundance of growth forms for fleshy fruits,
dry fruits often dispersed by wind (capsule, samara, and achene), and follicle and nut in the Australian Tropical Forest, Prairies and Plains of Central
North America, and Indian Desert, respectively. Tr, tree; S/SmT, shrub and small tree; H, herb; V/M, vine and mistletoe.


Journal of Systematics and Evolution Vol. 46 No. 3 2008 404
explosive) dispersal methods that are successful in
understory environments as well as open environ-
ments. This may in part explain the abundance of
capsules in the closed tropical forests of Australia. In
addition, many epiphytes have access to wind currents
in tropical forests and can therefore rely on wind
dispersal. Better knowledge of dispersal mechanisms
would make surveys more accurate since it may serve
as the source of selection pressure on fruit type and
may reveal more information on the evolutionary
patterns and significance of fruit types in various
environments.
The combined phylogenetic and ecological
analyses will begin to address the functional question
of whether the developmental constraint on fruit types
is weak enough to allow quick evolution of new fruit
types in response to natural selection. Obviously, this
question can also be addressed from developmental
genetic approaches, which has lagged much behind
floral evolutionary developmental studies. Neverthe-
less, such studies could be readily launched and
considerably accelerated with the availability of
genome sequences, functional genomic tools, and
growing knowledge of genes controlling fruit devel-
opment in model organisms (Liljegren et al., 2004;
Tanksley, 2004; Kellogg, 2006).
References
Bhandari MM. 1978. Flora of the Indian desert. India: United
Book Traders.
Bolmgren K, Eriksson O. 2005. Fleshy fruits—origins, niche
shifts, and diversification. Oikos 109: 255–272.
Clark CJ, Poulsen JR, Parker VT. 2001. The role of arboreal
seed dispersal groups on the seed rain of a lowland
tropical forest. Biotropica 33: 606–620.
Cooper W. 2004. Fruits of the Australian tropical rainforest.
Australia: Nokomis Editions.
Herrera CM. 1989. Seed Dispersal by Animals: A role in
angiosperm diversification? The American Naturalist 133:
309–322.
Eriksson O, Friis EM, Lofgren P. 2000a. Seed size, fruit size,
and dispersal systems in angiosperms from the early
Cretaceous to the late Tertiary. The American Naturalist
156: 47–58.
Eriksson O, Friis EM, Pedersen KR, Crane PR. 2000b. Seed
size and dispersal systems of early Cretaceous
angiosperms from Famalicao, Portugal. Plant Science 161:
319–329.
Heywood, VH. 1993. Flowering plants of the world. Updated
Edition. New York: Oxford University Press.
Judd WS, Campbell CS, Kellogg EA, Stevens PE, Donoghue
MJ. 2002. Plant systematics: A phylogenetic approach.
Sunderland, MA: Sinauer Associates.
Kellogg EA. 2006. Progress and challenges in studies of the
evolution of development. Journal of Experimental Botany
57: 3505–3516.
Knapp S. 2002. Tobacco to tomatoes: a phylogenetic
perspective on fruit diversity in the Solanaceae. Journal of
Experimental Botany 53: 2001–2022.
Kubitzki K, Rohwer JG, Bittrich V. 1993. The families and
genera of vascular plants. Heidelberg, Berlin: Springer-
Verlag.
Liljegren S, Roeder A, Kempin S, Gremski K, Ostergaard L,
Guimil S, Reyes D, Yanofsky M. 2004. Control of fruit
patterning in Arabidopsis by INDEHISCENT. Cell 116:
843–853.
Morgan DR, Soltis DE, Robertson KR. 1994. Systematic and
evolutionary implications of rbcL sequence variation in
Rosaceae. American Journal of Botany 81: 890–903.
Motley TJ, Wurdack KJ, Delprete PG. 2005. Molecular
systematics of the Catesbaeeae-Chiococceae complex
(Rubiaceae): flower and fruit evolution and biogeographic
implications. American Journal of Botany 92: 316–329.
Rydberg PA. 1932. Flora of the Prairies and Plains of Central
North America. New York: New York Botanical Garden.
Smith JF. 2000. A phylogenetic analysis of tribes Beslerieae
and Napeantheae (Gesneriaceae) and evolution of fruit
types: Parsimony and maximum likelihood analyses of
ndhF sequences. Systematic Botany 25: 72–81.
Tanksley SD. 2004. The genetic, developmental, and molecular
bases of fruit size and shape variation in tomato. Plant Cell
16: S181–189.
Tiffney BH. 2004. Vertebrate dispersal of seed plants through
time. Annual Review of Ecology, Evolution and
Systematics 35: 1–29.
van der Pijl L. 1982. Principles of dispersal in higher plants.
New York: Springer-Verlag.
Willson MF, Traveset A. 2000. The ecology of seed dispersal.
In: Fenner M ed. Seeds: The ecology of regeneration in
plant communities. 2nd ed. Wallingford: CAB Inter-
national. 85–110.
Zjhra ML, Sytsma KJ, Olmstead RG. 2004. Delimitation of
Malagasy tribe Coleeae and implications for fruit
evolution in Bignoniaceae inferred from a chloroplast
DNA phylogeny. Plant Systematics and Evolution 245:
55–67.